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Inland Fishes of Massachusetts 



Inland Fishes 
of Massachusetts 

Karsten E. Hartel, David B. Halliwell, and Alan E. Launer 

Illustrated by Laszlo Meszoly 



A 



Massachusetts Audubon Society 

Natural History of New England Series I Christopher W. Leahy, General Editor 



mcz 

8RARV 

OCT 1 6 2002 

H fi 

UNIVERSITY.' 

Copyright © 2002 by Karsten E. Hartel, David B. Halliwell, 
and Alan E. Launer. 

Published by the Massachusetts Audubon Society, 
208 South Great Road, Lincoln, MA 01773 

All rights reserved. No part of this book may be reproduced or 
transmitted in any form by any means, electronic or mechanical, 
including photocopying and recording, or by any information or 
retrieval system, without permission in writing from the 
Massachusetts Audubon Society, publisher. 

Library of Congress Cataloging-in-Publication Data 

Hartel, Karsten E. 

Inland fishes of Massachusetts / Karsten E. Hartel, David B. 
Halliwell, and Alan E. Launer ; illustrated by Laszlo Mezoly. 

p. cm. — (Natural history of New England series) 
Includes bibliographical references (p. ). 

ISBN 0-932691-28-5 (hard cover) 

1. Freshwater fishes — Massachusetts. I. Halliwell, David B. 
II. Launer, Alan E. III. Massachusetts Audubon Society. 
IV. Title. V. Series. 

QL628.M4 H37 2002 

597'.09744— dc21 

2002007490 



This book is written for those who care about and enjoy 
Massachusetts wetlands and the resident aquatic animals and plants. 

It is particularly dedicated to the following individuals 
who have contributed to our understanding of local fishes, 
aquatic ecology, and fisheries management. 

THOMAS J. ANDREWS, BRITTON C. McCABE, PETER H. OATIS, ROGER J. REED 



Contents 



Preface ix 
Acknowledgments xi 

Introduction 1 

Ichthyology in Massachusetts 8 

Conserving and Enjoying Fishes and the Aquatic Environment 2 1 

The Land, Water, and Fishes of Massachusetts 27 

How to Use This Book 39 



Identification Keys and Species Accounts 49 

Key to the Families and Monotypic Species of Massachusetts Inland 
Fishes 51 



Lampreys 60 

Lampetra, Petromyzon 

Sturgeons 66 

Acipenser 



Bowfins 

Amia 



71 



Freshwater eels 

Anguilla 

Herrings 78 

Alosa, Dorosoma 



74 



Anchovies 

Anchoa 



89 



Carp and Minnows 92 
Carassius, Couesius, Cyprinus, 
Hybognathus, Luxilus, Notemigonus, 
Notropis, Phoxinus, Pimephales, 
Rhinichthys, Scardinius, Semotilus 

Suckers 135 

Catostomus, Erimyzon 

Bullhead catfishes 143 

Ameiurus, Ictalurus 

Pike and pickerels 157 
Esox 

Mudminnows 165 

Umbra 



Vll 



Smelt 168 

Osmerus 

Salmon, chars, and trout 171 

Oncorhynchus, Salmo, Salvelinus 

Trout-perch 186 

Percopsis 

Cods 189 

Lota, Microgadus 

Needlefishes 195 

Strongylura 

Killifishes and pupfishes 198 

Cyprinodon, Fundulus, Lucania 

Silversides 211 

Menidia 

Mullets 216 

Mugil 

Sticklebacks 219 

Apeltes, Gasterosteus, Pungitius 

Literature Cited 283 



Pipefishes 229 

Syngnathus 

Sculpins 231 

Cottus 

Striped basses 235 

Morone 

Sunfishes and black basses 241 

Ambloplites, Enneacanthus, Lepomis, 
Micropterus, Pomoxis 

Perches and darters 266 

Etheostoma, Perca, Stizostedion 

Jacks 277 
Caranx 

American soles 279 

Trinectes 



Appendices 

1 . Indexed References 315 

2 . Distribution Table of Massachusetts Inland Fishes 317 

3. Glossary 321 

Taxonomic Index 325 

Color illustrations follow page 208 



viii Inland Fishes of Massachusetts 



Preface 



Why a new book on Massachusetts fishes? 

For some people, the answer to this question may be linked to the simple 
pleasure of observation. The ancient, yearly spectacle of the spawning 
migration of fishes into our coastal streams — though sadly dwindled — 
continues to inspire wonder for anyone who makes the effort to follow 
them on their aquatic journeys. 

For other people, a fascination with fishes is linked to the recreational 
pursuit of species that offer sustenance as well as the opportunity to recon- 
nect with the natural world, a need felt more acutely in our largely urban 
and technological culture. Still others recognize that protecting the diver- 
sity of life on this planet is essential to the health of ecosystems upon which 
all of us depend. Fishes are a vital part of the local ecosystem and food web 
because they eat smaller animals, plants, and fishes, and, in turn, they serve 
as food for many other creatures. 

The diversity of native freshwater fishes in Massachusetts is modest due 
to natural limiting factors such as geology, climate, and water chemistry. To 
these natural pressures on our native fauna, we have added stresses directly 
related to intensive, widespread human settlement. These include the ex- 
cessive use of surface water and groundwater, the clearing of much of the 
land's mature vegetation — predominantly temperate forests in Massachu- 
setts — and the conversion of forested areas to intensive agricultural use 
beginning early in the colonial period. Subsequently, industrialization ex- 
ploited the region's waterways for transportation, energy production, and 
the disposal of various, often toxic, wastes. Finally, we have introduced a 
variety of exotic fish species, either to improve sportfishing opportunities 
or through the careless release of live bait. As a result of all these human in- 
terventions, populations of some native inland species of fishes in Massa- 
chusetts have been reduced; and today introduced species comprise 45 per- 
cent of the state's primary freshwater ichthyo fauna. 



IX 



We live at a point in the earth's history when extinction of the planet's 
flora and fauna is proceeding at an unprecedented and alarming rate. Any 
efforts to slow and eventually stop this trend must be predicated upon 
knowledge of key facts: Which species occur in which habitats and water- 
sheds? What aspects of their life histories are critical to their survival? Which 
species are threatened with obliteration and why? 

The publication of Inland Fishes of Massachusetts gives us, for the first 
time, the answers to many crucial questions and makes a splendid contri- 
bution to our understanding of the Commonwealth's natural history. Here 
are refreshingly decipherable keys to identification — accompanied by pre- 
cise line drawings and color photographs; comprehensive species accounts 
with sections on identification, natural history, distribution, and abundance; 
summaries of Massachusetts ichthyology, fisheries, ecology, conservation, 
and fish watching; and an extensive bibliography. 

Inland Fishes of Massachusetts establishes the baseline against which all 
future studies of the freshwater fishes of this state will be measured. It is an 
indispensable addition to the library of all New England naturalists and 
sportfishers. 

Gary R. Clayton 

Vice President for Programs 

Massachusetts Audubon Society 



x Inland Fishes of Massachusetts 



Acknowledgments 



In a project that has spanned over two decades, there are many people to 
acknowledge and thank — we hope only that no one has been forgotten. Spe- 
cial thanks go to our wives, JoAnne Hartel, Susan Davies, and Elaine Launer. 
Many friends and colleagues, including J. Craddock, M. Estes, L. Kaufman, 
L. La Pointe, K. Liem, and M. Stiassny, have supported us through parts of 
the project. Douglas G. Smith, University of Massachusetts, Amherst, was 
originally a co-author of this effort but stepped aside to put more time into 
his studies of local invertebrates. He assisted at many levels and on many 
facets of this book and deserves our special thanks. Thomas J. Andrews, for- 
merly of the University of Massachusetts, Amherst, also was on the original 
team that shared the thought of an authoritative state checklist. The fish 
collections at the university stand as evidence of his efforts. 

Reviewers Dick Backus, Gary Clayton, Betsy Colburn, Bob Daniels, JoAnne 
Hartel, Todd Richards, Fritz Rohde, and Jim Williams read the complete 
manuscript at various stages and offered valuable assistance. We are grate- 
ful to the following for their comments on various sections: Ken Able, Joe 
Bergin, Jim Cardoza, Gary Clayton, Bruce Collette, Ed Crossman, Bill Eastes, 
David Etnier, Bill Fink, Arnie Howe, Bob Jenkins, Dick Keller, Bill Kreuger, 
Boyd Kynard, John Lundberg, Doug Markle, Amy McCune, Tom Monroe, 
Steve Murawski, Joe Nelson, Larry Page, Lynne Parenti, Fritz Rohde, Mike 
Ross, Doug Smith, and Melanie Stiassny. 

Artwork and maps We are grateful for the fine artwork of Laszlo Meszoly, 
who drew all of the fish drawings from photographs, specimens, and many 
references, including Bigelow and Schroeder (1953); Jenkins and Burkhead 
(1993); Pflieger (1975); Scott and Crossman (1973); and Trautman (1981). 
The base map used in this book was drawn and transferred to mylar by the 
Clark University Cartography and Information Graphics Service through 
the efforts of Anne Gibson. 



XI 



Records and museum specimens For access to museum specimens and 
records we are most grateful to T. Andrews, A. Richmond, and D. Smith 
(University of Massachusetts, Amherst); R. Bailey, W. Fink, and D. Nelson 
(University of Michigan); E. Bohlke and W. Saul (Academy of Natural Sci- 
ences, Philadelphia); J. Hoff (Southeastern Massachusetts University); 
J. Humphries and A. McCune (Cornell University); S. Jewett, J. Williams, 
and R. Vari (National Museum of Natural History, Smithsonian Institution); 
G. Jones (Northeastern University); G. Bond (Fitchburg State College); 
T. Graham (Framingham State College); W. Kenney (Springfield Science 
Museum); J. Eastman (Brown University); and M. Chandler and L. Kaufman 
(New England Aquarium). T. Whittier (US Environmental Protection Pro- 
gram) facilitated deposit of Emap specimens at the Museum of Compara- 
tive Zoology (MCZ). The Massachusetts Division of Fisheries and Wildlife 
and the University of Michigan donated Massachusetts specimens to the 
MCZ collection. 

Information and other favors A. Coleman and R. Jenkins offered us advice 
on photography. Information on local fishes was received from R. Schmidt 
and W. Kenney. W. Smith-Vaniz and V. Vladykov confirmed the identifica- 
tion of selected specimens. A.J. Screpetis supplied information on rivers 
and ponds. 

Massachusetts Division of Fisheries and Wildlife (MDFW) The Division 
supported the stream surveys of D. Halliwell between 1977 and 1990, and 
MDFW district fisheries supervisors, in particular Lee McLaughlin, and 
their field crews were instrumental in completing this work. Fisheries staff 
at the Westborough Field Headquarters, especially J. Bergin, B. Eastes, 
R. Hartley, D. Keller, and J. O'Leary, contributed in many ways. The late 
Peter Oatis allowed D. Halliwell to spend time on various aspects of this 
project and provided funds to make a number of the illustrations for this 
book. Updated information on fish occurrences was received from J. Bergin, 
T. Richards, and K. Simmons. R. Arini brought several new catfish records 
to our attention. 

Museum of Comparative Zoology (MCZ) Enormous thanks go to Karel F. 
Liem, Curator of Ichthyology, for unflinching support of this project since 
its inception. Almost all of the local field work out of the MCZ was supported 
out of our own pockets and succeeded only through the volunteer weekend 

xii Inland Fishes of Massachusetts 



efforts of M. Buckley, C. Gougeon, J. Jensen, G. Lauder, Jr., S. Norton, J. Ro- 
sado, F. Ross, and E. Wu. The MCZ Ichthyology Department supported cata- 
loguing of the field collections or donations of Massachusetts specimens 
from R. Gibbs and W. Kreuger (Boston University), K. Hartel, A. Launer, 
D.G. Smith, and the MDFW. The National Science Foundation supported 
the development and implementation of the MUSE database that expedited 
record keeping for this project. K. Blake, M. Buckley, J. Bush, C. Gougeon, 
J. Kelly, M. Stein, and D. Triant sorted and catalogued many of the speci- 
mens. A. Launer was supported under the Harvard graduate program in 
Organismic and Evolutionary Biology during part of his work on this book. 
P. Mclntyre assisted in the design of tables and graphs. 

Massachusetts Audubon Society Special thanks are due to Vanessa Rule 
and Mary Hopkins for coordinating the many complex aspects of this proj- 
ect through various stages of production, and to Denise Bergman for proof- 
reading in its final stages. 



Acknowledgments xiii 



Inland Fishes of Massachusetts 



Introduction 



This is the first detailed guide to Massachusetts freshwater fishes and their 
distribution. Its primary aim is to present updated information on the iden- 
tification and distribution of these fishes to both the beginner and the pro- 
fessional. In addition, we have included brief outlines about the natural 
history and status of the fauna. This book is designed to meet the needs of 
anglers, aquatic resource managers, conservation groups, educators, natu- 
ralists, and the general public. We hope that the information presented 
here will make the reader more aware of aquatic resources and the natural 
heritage of Massachusetts. 

Fishes have played an important role in Massachusetts' history. The Na- 
tive Americans used them as food and fertilizer, and the first European ex- 
plorers marveled at their abundance. The large number of fishes attracted 
numerous settlers to New England. Early colonists found the marine waters 
full of cod, herrings, and flatfishes and the estuaries and rivers rich in shad, 
sturgeons, and Striped Bass. Even the "lowly" Sea Lamprey supported a 
local fishery during this era. 

When the colonists wrote of the abundance of fishes they were referring 
to the great quantities of a few, largely marine, species and not to the num- 
bers of different kinds of fishes. Native freshwater fishes in Massachusetts 
are a naturally depauperate fauna. The low number of native species is due 
primarily to the glaciers that covered all of Massachusetts as recently as 
14,000 years ago. The intervening years are a short time in a geological sense, 
and only a few species found their way into the deglaciated areas. For ex- 
ample, 50 or so native species are known from New England, whereas nearly 
300 are known from the Appalachian region. 

Far beyond their obvious value to humans as a source of food and recre- 
ation, fishes are vital components of aquatic ecosystems. Fishes both eat 
and are eaten in a food chain where algae and invertebrates are converted 
into food for fishes and other animals, including humans. About one-third 
of all bird species found in Massachusetts eat fishes at some time during 

Introduction 1 



their lives. In fact, fishes are a life- sustaining necessity for some birds, in- 
cluding loons, grebes, cormorants, herons, mergansers, Bald Eagles, Os- 
preys, and terns. Fishes are also directly important to some mammals, rep- 
tiles, amphibians, and invertebrates: otters, mink, water shrews, turtles, 
snakes, diving beetles, and dragonfly larvae all eat fishes. In addition, the 
existence of most freshwater mussels would be impossible without fish be- 
cause larval mussels spend the first part of their lives attached to the gills 
of living fishes. 

The interwoven relationship of water, fishes, birds, other animals, and 
plants is complicated. It can be affected by atmospheric input from hun- 
dreds of miles away, downstream flow from other states, local construction, 
agricultural runoff, changes in groundwater, or the introduction of addi- 
tional species into the web. The delicate aquatic ecosystem must always be 
considered when studying fishes because their world is so closely related to 
and yet so very different from our own. 

Today, none of Massachusetts' freshwater fishes, except possibly eels and 
herrings, which are caught at sea, are harvested commercially. However, 
vast numbers of people enjoy the recreational value of fishing the state's 
waters. Between 1986 and 1995, approximately 270,000 fishing licenses were 
sold each year in Massachusetts. These fees generated 3.5 million dollars 
annually for the Massachusetts Division of Fisheries and Wildlife (Cardoza 
1997, pers. comm.). A high percentage of these funds is directed to game- 
fish related activities that often indirectly benefit the full aquatic ecosystem. 
In addition, a national survey (Anon. 1993) estimates that some 652,000 an- 
glers spent 454 million dollars while pursuing their sport in the marine and 
freshwater areas of Massachusetts in 1991. 

Outline of This Book 

This book focuses on Massachusetts freshwater fishes and does not cover 
the local marine species or fishes found outside of the state; nor does this 
guide deal in detailed accounts offish anatomy, physiology, or behavior. 
However, small amounts of what we consider interesting related informa- 
tion are scattered throughout the book, and we have provided a general ref- 
erences section that indexes topics beyond the scope of this work. 

It should be emphasized that this book and its identification keys are de- 
signed to be used with fishes found in Massachusetts freshwaters. Although 



Inland Fishes of Massachusetts 



they apply generally to most New England freshwater fishes, they will not 
work with some species from the Lake Champlain and Saint Lawrence 
drainages of northern Vermont, New Hampshire, and Maine. 

In addition to introductory material, this book is divided into three parts: 
illustrated keys for identification of families and species; an annotated syn- 
opsis of all families and species known from the state, with selected com- 
ments on their biology and conservation; and reference material. The refer- 
ence material includes literature cited, general fish references (Appendix 1), 
a table summarizing the distribution of each species by drainage areas (Ap- 
pendix 2), and a glossary (Appendix 3). 

Fishes in General 

Fishes may be best described as vertebrate animals that breathe through 
gills and have median fins with skeletal supports. However, many of the ap- 
proximately 25,000 species found worldwide lack one or more of the com- 
mon attributes often associated with fishes: many lack scales, others are 
blind, and some even breathe air. Fishes are truly remarkable in that they 
occur in a far wider variety of habitats than any other vertebrate group. They 
live in areas that range from high mountain lakes at 12,500 feet, to ocean 
canyons deeper than five miles. Certain species can survive the subfreezing 
Antarctic water by using a natural antifreeze in their blood, and others have 
adapted to African hot soda lakes at temperatures close to 120°F. Fishes 
break many general biological rules in various ways: some change their sex, 
die almost immediately after reproducing, catch food on land, produce an 
electric field for communication and prey capture, find their homestreams 
by smell or have all female species, and some fishes can even be said to 
"sing" or to "fly." In general, Massachusetts freshwater fishes are not quite 
so diverse in their habits, but each species is fascinating in its own right and 
has an interesting story to tell. 

Massachusetts Fishes — Who's Related to Whom 

Three major divisions of the vertebrates are most often called fishes: the 
jawless fishes (lampreys and hagfishes); the cartilaginous fishes (sharks, 
skates, and rays); and the bony, ray-finned fishes (minnows, trout, bass, 
and many others). The relationships and classification of most of the major 



Introduction 



Table 1 A classification of the families of freshwater fishes found in Massa- 
chusetts (modified from Nelson 1994, Stiassny et al. 1996). 

JAWLESS FISHES (formerly AGNATH A) 

Cephalaspidomorphi Petromyzontiformes 

Petromyzontidae (lampreys) 
GNATHOSTOMATA 

Actinopterygii Acipenseriformes 

Acipenseridae (sturgeons) 
Amiiformes 

Amiidae (bowfins) 



Teleostei 

Elopomorpha 

Anguilliformes 

Anguillidae (freshwater eels) 
Otocephala 

Clupeomorpha 

Clupeiformes 

Clupeidae (herrings) 
Engraulidae (anchovies) 
Ostariophysi 

Cypriniformes 

Cyprinidae (minnows) 
Catostomidae (suckers) 
Siluriformes 

Ictaluridae (bullhead catfishes) 
Euteleostei 

Protocanthopterygii 

Salmoniformes 

Salmonidae (salmon) 
Osmeridae (smelt) 
Neognathi 

Esociformes 

Esocidae (pikes and pickerels) 
Umbridae (mudminnows) 
Paracanthopterygii 

Gadiformes 

Gadidae (codfishes) 
Percopsiformes 

Percopsidae (trout-perches) 
Acanthopterygii 

Mugilomorpha 

Mugiliformes 

Mugilidae (mullets) 
Atherinomorpha 

Atheriniformes 

Atherinopsidae (silversides) 
Beloniformes 

Belonidae (needlefishes) 



Inland Fishes of Massachusetts 



Table 1 (continued). 



Cyprinodontiformes 

Fundulidae (killifishes) 
Cyprinodontidae (pupfishes) 
Percomorpha 

Gasterosteiformes 

Gasterosteidae (sticklebacks) 

Syngnathiformes 

Syngnathidae (pipefishes) 

Scorpaeniformes 

Cottidae (sculpins) 

Perciformes 

Moronidae (striped basses) 
Centrarchidae (sunfishes) 
Percidae (perches) 
Carangidae (jacks) 

Pleuronectiformes 

Achiridae (American soles) 



groups of Massachusetts freshwater fishes are shown in Table 1. In variety, 
our local fishes range from the most "primitive" lampreys to the most "ad- 
vanced" bony fishes, the teleosts. 

Within these three divisions, Massachusetts fishes are further split into 
classes, orders (names ending in "iformes"), and families (names ending in 
"idae"). Orders contain closely related families, and families contain related 
genera and species. Thus, each species fits into a multi-tiered classification. 
Table 1 shows all of the families of fishes found in Massachusetts in a hier- 
archical format. Information from the study of anatomy, life history, and 
biochemistry allows scientists to link or separate groups of animals. Much 
has been learned over the last two decades about the interrelationships of 
fishes, but there are a few groups in which distantly related fishes are still 
lumped together into unnatural (paraphyletic or polyphyletic) groups. The 
study of relationships, called "systematics," places animals in a framework 
that allows comparisons between and among groups and, therefore, is cru- 
cial to the understanding of variations in fish behavior, physiology, ecology, 
and management. 

The most basal of Massachusetts' fishes are the jawless lampreys, which 
have survived hundreds of millions of years with little change. Although 



Introduction 



seemingly primitive, they are well adapted to their habitat and behavior. 
The lampreys were at one time placed in the class Agnatha, but that assem- 
blage has recently been called into question as lampreys are probably more 
closely related to jawed fishes than to the jawless hagfishes. All other verte- 
brates, from sharks to mammals, belong to the jawed Gnathostomata. 

The jawed fishes are divided into the cartilaginous sharks, skates, and 
rays, and the bony fishes. The sharklike forms have individual gill slits and 
lack a bony skeleton. While none of these fishes occur in Massachusetts 
freshwaters, some sharks have been found as far up the Mississippi River as 
Illinois. The bony fish group contains the ray-finned fishes (Actinopterygii) 
and the lobe-limbed vertebrates (Sarcopterygii). Many extinct fishes, along 
with the living lungfish (Dipnoi), and all amphibians, reptiles, birds, and 
mammals belong to the lobe-limbed group. 

Sturgeons and the Bowfin are Massachusetts' representatives of two an- 
cient groups of primitive ray-finned fishes that date back 300 million years 
and have few surviving members. All other Massachusetts ray-finned fishes 
are members of the more advanced Teleostei, a diverse group with some 
20,000 species found in virtually every aquatic habitat around the world. 
Teleosts have posterior vertebrae that expand into plates that give support 
to a symmetrical tail or caudal fin. 

Massachusetts teleosts can be arranged into six subgroups: the eels 
(Elopomorpha); the herrings (Clupeomorpha); the minnows and relatives 
(Ostariophysi); the salmonlike fishes (Protocanthopterygii); the pike, pick- 
erels, and relatives (Neognathi); the codlike fishes placed in the paraphy- 
letic Paracanthopterygii; and the more advanced teleosts (Acanthopterygii) . 
The fishes in the eel group are combined because they have a specialized, 
wafer-thin larva, the leptocephalus. The herringlike fishes, which include 
herrings and anchovies, all have a stethoscopelike connection between the 
swim bladder and the ear. The minnows, suckers, and catfishes are placed 
in the Ostariophysi because of the development of the anterior vertebra 
into a specialized organ called the "Weberian apparatus." 

The last major teleost subgroup, the Acanthopterygii, is united based on 
the position of a muscle associated with the upper set of pharyngeal jaws. 
Acanthopterygian fishes of Massachusetts include the mullets (Mugilo- 
morpha), the silversides, killifishes, pupfishes, and needlefishes (Atheri- 
nomorpha), and the advanced spiny- rayed fishes (Percomorpha). The 
spiny-rayed fishes encompass the widest array offish families: the stickle- 



6 Inland Fishes of Massachusetts 



backs, the sculpins, the perchlike fishes, and the specialized flatfishes 
(Pleuronectiformes) . 

references. Lauder and Liem 1983; Moser et al. 1984; Johnson 1992; 
Burr and Mayden 1992; Stiassny et al. 1996 (relationships); Nelson 1994; 
Eschmeyer, ed. 1998 (systematic list of all fish groups); Wiley 1981; 
Mayden and Wiley 1992 (systematic practice and theory). 



Introduction 7 



Ichthyology in Massachusetts 



"There are in the rivers, and ponds, very excellent Trouts, Carpes, Breames, 
Pikes, Roches, Perches, Tenches, Eeles, and other fishes, such as England doth 
afford, and as good, for variety; yea many of them much better..." Thomas 
Morton writing of the Massachusetts Bay area in the 1630s (Morton 1972). 

The Study of Fishes 

Although Massachusetts had been explored by Europeans as early as 1602, 
little ichthyological information was published until the early 1800s. The 
first in-depth freshwater fish survey of Massachusetts, including only the 
western portion of the state, was not undertaken until 1940. It is unfortu- 
nate that surveys were not made in the earlier years because many ques- 
tions will never be answered about the former distribution and abundance 
of Massachusetts fishes. 

Prehistoric Records The first records of postglacial Massachusetts fishes 
come from sparse remains of fishes found at Native American archaeologi- 
cal sites that date from 5,000 years ago to the early 1600s. The compara- 
tively small amount of New England archaeological material is probably 
due to the methods of food processing or its transportation from site to site 
and the poor quality of the preservation offish bones in New England's cli- 
mate and acidic soils. A recent review offish remains at archaeological sites 
in New England (Carlson 1988) lists 27 marine species, 6 anadromous spe- 
cies, and 4 groups of freshwater fishes (minnows, catfishes, sunfishes, and 
perches). In addition, Huntington (1982) found remains of Brook Trout at 
a site in Marlborough, and Fallfish and Chain Pickerel have been recently 
identified from an important shell heap along the Sudbury River (Smith 
1940, Largy 1995). 

Nearly all local fishes might have been used by the Native Americans, but 
it seems odd that more remains of freshwater fishes have not been discov- 



8 Inland Fishes of Massachusetts 



ered at inland archaeological sites. Remains of anadromous species have 
frequently been encountered at both inland and estuarine sites in Massa- 
chusetts; bones from Atlantic Tomcod, river herrings, and American Eels 
have been found at a site in Marlborough (Huntington 1982), American 
Shad and river herrings at Turners Falls, and sturgeons along the Merrimack 
River. William Wood (Vaughan 1977) reports that the first Europeans saw 
the Native Americans fishing for sturgeon with strong nets during the day 
and with torches and spears at night. 

Atlantic Salmon have been logically considered a seasonal mainstay in 
New England by many historians but, surprisingly, Atlantic Salmon remains 
have not been found at any Massachusetts archaeological sites and have 
been positively reported at only one site in Maine. A recent study by Carlson 
(1988 : 65) states "archaeological faunal evidence for fishing in New England 
does not suggest a heavy utilization of Atlantic Salmon [Salmo salar] in ei- 
ther coastal, estuarine, or riverine settings..." and "...all available evidence 
indicates that salmon was an extremely minor component of the prehistoric 
resource base," but the results of this study may be biased by centuries of 
New England climate. 

The Early Years (1 600 to 1830) The first records of the fishes of New En- 
gland come from the journals and notes of the early adventurers. Most of 
these reports mention only species of economic importance or of curious 
nature; almost all fail to mention freshwater fishes. The earliest accounts, 
from Bartholomew Gosnold's 1602 voyage to Cape Cod and Buzzards Bay, 
mention eight species of marine fishes and comment on their quality and 
abundance (Archer 1843, Brereton 1906). John Smith's accounts briefly 
mention sturgeons, perches, and eels (Smith 1986). The first actual list of 
species by John Josselyn in 1672 contains about 46 species and concen- 
trates primarily on marine food fishes, but it also includes species from as 
far south as Virginia (Lindholdt 1988). 

Although the early reports offer an incomplete picture of the fishes of 
Massachusetts, one feature is fairly well documented — fishes were once 
far more abundant than they are today. Almost all early accounts mention 
great quantities of herrings, sturgeons, Striped Bass, and Atlantic Salmon. 
Some of the accounts seem believable, while others contain more fanciful 
descriptions. Wood writes in 1634 (Vaughan 1977) that "sturgeons be all 
over the country, but the best catching of them is upon the shoals of Cape 
Cod and in the river of Merrimack, where much is taken, pickled and 



Ichthyology in Massachusetts 9 



brought to England. Some of these be twelve, fourteen, eighteen foot long." 
Alewives are "in such multitudes as is almost incredible, pressing up in such 
shallow waters as will scarce permit them to swim, having likewise such 
longing desire after the fresh water ponds that no beatings with poles or 
forcive agitations by other devices will cause them to return to the sea till 
they have cast their spawn." Wood further states that "below this fall of 
waters [Charles River at Watertown] they [the inhabitants of the town] take 
great stores of shads and alewives. In two tides they have gotten one hun- 
dred thousand...." Striped Bass were likewise described: "some be three 
and some be four foot long, some bigger, some lesser. At some tides a man 
[with hook and line] may catch a dozen or twenty in three hours." In the 
spring, Striped Bass were taken in rivers with nets, "sometimes two or three 
thousand at a set." 

In 1804, the first American scientific paper on New England fishes was 
written by William Dandridge Peck (1763-1822). This work, entitled De- 
scription of Four Remarkable Fishes taken near the Piscataqua in New 
Hampshire, did not describe freshwater species. However, Peck, who was 
appointed natural history professor at Harvard in 1805, did collect some 
freshwater fishes. The Harvard Museum of Comparative Zoology houses 
1 1 dried skins of New England freshwater fishes, collected by Peck between 
1790 and 1793, which have handwritten labels showing that many are from 
the Piscataqua River; however, one Rainbow Smelt is labeled from Boston. 
The other species include the Sea Lamprey, Alewife, White Perch, Pump- 
kinseed, and Yellow Perch. These dried preparations represent some of the 
oldest natural history material available from the United States. 

In 1816, Charles A. Lesueur (1778-1846), a French ichthyologist, visited 
Boston shortly after arriving in the United States and described a number 
of local species based on the material that he found. All these species, how- 
ever, had been described by earlier ichthyologists, especially by Samuel 
Latham Mitchill in his accounts of New York fishes. Mitchill and two other 
New Yorkers, Alexander Wilson and DeWitt Clinton, preceded Lesueur and 
all the early Massachusetts ichthyologists in first describing many of the 
East Coast species from their studies between 1814 and 1824. 

The Middle Years (1 830 to 1 900) The first lists of Massachusetts fishes 
were produced by a physician and one-term mayor of Boston, Jerome Van 
Crowningshield Smith (1800-1879; see Figure la) and included in Edward 
Hitchcock's 1833 and 1835 reports on the geology of Massachusetts. The 

1 Inland Fishes of Massachusetts 





Figure la. J.V.C. Smith (1800-1879) com- 
piled the first list of Massachusetts fishes. 
Courtesy Boston Public Library. 



lb. D.H. Storer (1804-1891) prepared 
two major works and many papers on 
Massachusetts fishes. Courtesy Boston 
Museum of Science. 



1835 report lists about 25 species of freshwater fishes that we can recognize 
based on the names that he used. In 1833, Smith had also produced a book 
called Natural History of the Fishes of Massachusetts, Embracing a Practical 
Essay on Angling, which was released again in 1843 with almost no revisions. 
Smith's work was severely criticized by his contemporary, D.H. Storer, and 
later by Theodore Gill in his history of the state's ichthyology (1904). Both 
Storer and Gill disapproved of Smith's fanciful accounts, his outdated tax- 
onomy, his inclusion of European fishes, the use of European names for 
many local fishes, and his unabashed publication of illustrations from un- 
credited sources. 

David Humphries Storer (1804-1891) (Garman 1891, Gifford 1964; see 
Figure lb) was perhaps the most important figure in Massachusetts ichthy- 
ology. Storer was a Boston physician who published a number of medical 
papers as well as more than 60 articles on ichthyology, herpetology, and 
conchology over 30 years. His first papers were presented at meetings of 
the Boston Society of Natural History (now the Boston Museum of Science), 
which he helped to found. Among his earliest papers were the critiques of 



Ichthyology in Massachusetts 1 1 



Jerome Van Crowningshield Smith's lists (1836) and his first history of Mas- 
sachusetts fishes. 

In 1837, Storer was commissioned to produce a report to the state com- 
missioners on the ichthyology and herpetology of Massachusetts. This en- 
deavor, financed by the Massachusetts legislature, was produced in 1839. 
Storer expanded this work in a series of six papers called A History of the 
Fishes of Massachusetts, published in the Memoirs of the American Acad- 
emy of Arts and Science between 1853 and 1867. The full series, published 
as one volume in 1867, contains almost 300 pages, 39 lithographed plates, 
and an appendix by Frederick Ward Putnam, an anthropologist and natural 
historian, that listed additional species. Storer's History is equivalent to the 
best of the state books of the period and was beautifully illustrated by Au- 
guste Sonrel (Blum 1993). Like Lesueur, Storer named a number of species 
that had already been described by earlier authors. Only two of his New 
England freshwater species are considered taxonomically valid today: the 
Tessellated Darter, described from the Connecticut River near Hartford, 
and the Northern Pipefish, described from Nahant. 

In 1859, Louis Agassiz (Lurie 1988, Winsor 1991) founded the Museum of 
Comparative Zoology at Harvard University and made major expeditions 
to Lake Superior and later to South America. His assistant Samuel Garman 
(1846-1927) (see Summers and Koob 1997) published a few papers on local 
fishes; but, in retrospect, it seems that the study of Massachusetts freshwa- 
ter fishes held no deep interest for Agassiz and his colleagues. Presumably, 
they concluded that Storer had covered Massachusetts fishes in sufficient 
detail. Fortunately, a number of interesting freshwater specimens from this 
period were deposited in Agassiz's museum, among them some of the only 
records of the Longnose Dace and Slimy Sculpin from the Merrimack River 
in Massachusetts. 

From the mid- 1800s to the early 1900s, only a few additional papers 
on Massachusetts fishes appeared. In 1858, Charles Girard described the 
Swamp Darter and the Banded Sunfish from specimens collected in Massa- 
chusetts. In 1879, G. Brown Goode and Tarleton H. Bean of the United States 
Fish Commission published A List of the Fishes of Essex County, including 
those of Massachusetts Bay. This work deals primarily with the state's marine 
fauna, but also lists the freshwater species of Essex County. More impor- 
tantly, Goode and Bean's report updated and corrected much of the confu- 
sion regarding the scientific and common names of local fishes. 



12 Inland Fishes of Massachusetts 



The Quiet Years (1900 to 1940) In 1908, William Converse Kendall produced 
a List of the Pisces in the Fauna of New England series published by the Bos- 
ton Society of Natural History. Kendall's work, mainly a literature survey, 
also cites many Massachusetts specimens from the collections of the Bos- 
ton Society of Natural History. Over the years, most of the Boston Society's 
fish specimens, including some cited by Kendall, have been transferred to 
the Museum of Comparative Zoology. Kendall produced two monographs 
on the salmonid fishes of New England (char 1914, salmon 1935) that cover 
taxonomy, natural history, and historical status. He also published papers 
on silversides (1902), catfishes (1910), and smelt (1926). 

During this period, Carl L. Hubbs (1894-1979), one of the most distin- 
guished ichthyologists of the 20th century, spent a year at the Museum of 
Comparative Zoology reviewing North American freshwater fishes. Hubbs 
and his wife, Laura, made a number offish collections around Massachu- 
setts that shed interesting light on the relative abundance of the fishes dur- 
ing the late 1920s. The Hubbs' specimens are now housed at the University 
of Michigan in Ann Arbor. 

In 1925, Dr. Henry Bryant Bigelow, a Harvard professor and founder of 
the Woods Hole Oceanographic Institution, and his colleague, W.W. Welsh, 
turned their attention to marine fishes and produced the first edition of 
Fishes of the Gulf of Maine. This monumental faunal work contains accounts 
of many of the diadromous and estuarine fishes that are sometimes found 
in Massachusetts freshwaters. Fishes of the Gulf of Maine was rewritten in 
1953 by the team of Dr. Bigelow and William C. Schroeder, who together 
wrote numerous papers on sharks and rays. 

The Recent Years (1940 to the present) The summer of 1940 marked a ma- 
jor turning point in knowledge of Massachusetts freshwater fishes. During 
that field season, Prof. Britton C. McCabe (1901-1968) (see Figure 2a) of 
Springfield College made 400 collections of fishes from sites in western Mas- 
sachusetts (Map 1). McCabe's fieldwork formed the basis of his doctoral 
thesis from Cornell University on the fishes of the streams of western Mas- 
sachusetts (McCabe 1942, 1943); this study was the first comprehensive fish 
survey in Massachusetts. McCabe became chairman of the Biology Depart- 
ment at Springfield College (1946-1963) and, during the summers between 
1944 and 1952, was an aquatic biologist for the Massachusetts Division of 
Fisheries and Wildlife. Prof. McCabe initiated a series of lake and pond sur- 



Ichthyology in Massachusetts 13 




Figure 2a. B.C. McCabe (1901-1968) 
conducted the first in-depth survey 
of the state's freshwaters. Courtesy 
Mrs. B.C. McCabe. 



2b. R.J. Reed (1929-1979) conducted 
research and taught at the University 
of Massachusetts. Courtesy Mrs. R.J. 
Reed. 



veys, from which he and his colleagues compiled six reports (McCabe 1948, 
1952, 1953; McCabe and Swartz 1952; Stroud 1955; Swartz 1944). 

Other colleges and universities began adding information in the mid- 
1900s. Prof. Thomas J. Andrews (University of Massachusetts, Amherst) 
made numerous collections all across Massachusetts, either by himself or 
with students, between 1948 and 1980. Andrews was the first to identify the 
Shortnose Sturgeon (Vladykov and Greeley 1963) and Bluntnose Minnow in 
Massachusetts. He avidly collected darters and studied their distribution 
and was the first to recognize the Swamp Darter on Martha's Vineyard. 

From 1958 to 1963, Dr. Robert H. Gibbs (1929-1988) taught at Boston 
University. With students, he made a number of collections, especially in 
eastern Massachusetts. Gibbs' Boston University fish collections, trans- 
ferred to the Museum of Comparative Zoology in 1978, contain records of 
some species, particularly the Bridle Shiner, from sites in eastern Massachu- 
setts where they no longer occur. 

In 1969, Paul S. Mugford (MDFW) produced the first modern guide to 
Massachusetts freshwater fishes, the Illustrated Manual of Massachusetts 
Freshwater Fish. This small book served the angling community for almost 
25 years, but now its information is out-of-date. 



14 Inland Fishes of Massachusetts 







Map 1 McCabe Data: localities surveyed by Britton 
McCabe 1940-1941. Open circles indicate sites where 
fishes were not found. 



Also associated with the universities are the Cooper- 
ative Fisheries and Wildlife Units, now under the United 
States Geological Survey. The Massachusetts Cooperative Fisheries Unit at 
the University of Massachusetts, Amherst, is a federal research station co- 
operating with the university and the state. Much of the work at Amherst 
has revolved around the study of the anadromous fishes of the Connecticut 
and Parker rivers. Dr. Roger J. Reed (1929-1979) (see Figure 2b), Coopera- 
tive Unit Leader at Amherst and one of the driving forces behind restoration 
of the state's anadromous fishes, was much broader in his research. Reed and 
his students actively studied a broad spectrum, including parasites, aquatic 
invertebrates, and nongamefish species. Almost half of Reed's publications 
are on such topics, including life histories of the Fallfish (1971), Blacknose 
Dace (Reed and Moulton 1973), and Tessellated Darter (Layzer and Reed 
1978) in Massachusetts and the Longnose Dace in Pennsylvania (1959). 

Over the last two decades, ichthyological research has continued. David 
Halliwell produced the updated A List of the Freshwater Fishes of Massachu- 
setts (1979 et seq.) and a Ph.D. thesis on Massachusetts streams and fish 
distribution (1989). During this time, Karsten Hartel at Harvard and Doug- 
las G. Smith at the University of Massachusetts, Amherst, began a concerted 
effort to collect fishes all across the state. Hartel and Halliwell shortly joined 
forces with the long-term goal to produce this book. Lists of exotic or intro- 
duced Massachusetts fishes were produced by Hartel (1992) and Cardoza 
etal. (1993). 

Anadromous fish work has expanded at the new S.O. Conte Anadromous 
Fish Research Lab at Turners Falls, which went into operation under the 
US Fish and Wildlife Service in 1990. Drs. Boyd Kynard, Henry Booke (now 






Ichthyology in Massachusetts 15 



retired), and Steve Rideout continue to study many species at that site. 
Active programs in ichthyology or fisheries biology exist at Southeastern 
Massachusetts University, the University of Massachusetts (Amherst and 
Boston), Boston University, and Harvard University. The New England 
Aquarium, Boston, through its research department, has also been instru- 
mental in producing critical studies of the state's aquatic fauna and flora. 

Fisheries, Past and Present 

Fisheries Management For centuries, people have increased the yield of 
harvestable fishes by capturing, holding, spawning, rearing, and otherwise 
manipulating them, by modifying their environment, and by protecting 
them from overharvesting. Massachusetts was the earliest state to produce 
formal fisheries laws and regulations. Colonial acts were established as early 
as 1709 to control the erection of weirs or other obstacles that prevented 
the passage of fishes. Seine, hook and line, and night fishing in specific 
bodies of water were regulated by the turn of the 19th century. By the mid- 
1800s, Massachusetts had appointed commissioners to examine the status 
of fisheries (1855), artificial propagation of fishes (1856), and dams and bar- 
riers (1865). In 1869, the state set up a permanent Commission on Inland 
Fisheries, with each of its three members serving five-year terms. 

During these early years, the state attempted the first official fish stock- 
ings. Rainbow Smelt were collected and transplanted to Boston's Jamaica 
Pond during the 1760s (Storer 1840). Black basses were introduced by 1850, 
American Shad fry were released into the Concord River in 1868, and the 
first Massachusetts hatchery at Agawam produced almost 40,000 salmon, 
trout, whitefishes, and chars between 1868 and 1870. 

As declines in anadromous fishes and sea fisheries were noted (Lyman 
1871), stocking continued at a fast pace to try to supplement or reestablish 
the native salmonid fishery. Hatcheries were established in the towns of 
Winchester (1870), Sutton (1902), Sharon, Norfolk, and Wilbraham (1912), 
and Sandwich (1914). 

Commercial fishing of inland waters stopped long before the turn of the 
century, and as recreational fisheries developed, the state instituted its first 
fishing licenses. In 1919, anyone fishing stocked waters was required to have 
a license, and, by 1930, a license was required to fish all inland waters. 

The Massachusetts Division of Fisheries and Wildlife (MDFW) is the 
agency that directly oversees the freshwater fisheries of Massachusetts. Its 

1 6 Inland Fishes of Massachusetts 



history can t>e traced back to the commissioners of the 1850s. Through the 
years, its primary concerns have often been with enhancement of recre- 
ational fisheries, but, since the late 1970s, this agency has gradually become 
more involved with the overall aquatic ecosystem. However, over the past 
several decades, research and protection of both game and nongame fishes 
are becoming balanced. The division's Endangered Species and Natural 
Heritage Program, the primary state agency responsible for rarer fishes, is 
almost completely supported by donations from the citizens of Massachu- 
setts through an income tax write-off system. 

Success at the Fishways One of the most dramatic and noticeable declines 
of New England fishes before the loss of marine ground fisheries late in the 
20th century (Fogarty and Murawski 1998) was caused by dam construc- 
tion, which eliminated thousands of miles of stream habitat for anadromous 
fishes. As settlements were established, canals and then dams were built to 
aid commerce and manufacturing. By 1798, a dam was in place in Massa- 
chusetts at Turners Falls in Montague on the Connecticut River. This dam 
was followed by structures at Holyoke on the Connecticut (1849) and Law- 
rence and Lowell (1847) on the Merrimack. These dams limited migratory 
runs to less than 90 miles on the Connecticut and about 30 miles on the 
Merrimack, thus eliminating vast areas of the river basins for reproduction 
and juvenile growth of such important migratory species as sturgeon, At- 
lantic Salmon, American Shad, Alewife, and Blueback Herring. 

By 1865, state-appointed commissions were examining the problem, 
and the Supreme Court ruled that the owners of the dams were responsible 
for installing and maintaining fishways. The first fishways were put in place 
by the early 1890s, but the long series of attempts to construct or improve 
fish passage, which lasted until almost 1950, was largely ineffective. Stock- 
ing was also attempted during these years but was negated by the fact that 
returning adult fishes could not swim upriver. By 1949, court orders and 
new technology came into play when the Holyoke Power Company was 
mandated to build fish passage facilities around their new power station 
at Holyoke. The first designs met with little success until 1955, when fed- 
eral fisheries personnel and the power company built the first successful 
upstream passage facility in the Northeast (Moffitt et al. 1982). This eleva- 
tor system is a large box or hopper into which fishes are attracted by cur- 
rent, captured, lifted to the height of the dam, and released into the upper 
impoundment. The operation of this facility opened 36 miles of river be- 

Ichthyology in Massachusetts 17 



tween Holyoke and Turners Falls to diadromous fishes for the first time in 
100 years. 

All of the major fishways in Massachusetts, at Westfield, Holyoke, Turn- 
ers Falls, Lowell, and Lawrence, are now operational. In addition, the estab- 
lishment of the S.O. Conti Anadromous Fish Laboratory at Turners Falls, 
which has a large flume area to test designs of new fishways, will help to 
improve the designs. Operation of the fishways, stocking programs, and re- 
lated research are expensive and would not have been possible, and will 
not continue, without town, state, and federal agencies cooperating with 
conservation agencies and the large power companies. 

The fishways have worked for many of the anadromous species. For ex- 
ample, on the Connecticut River over 8 million American Shad and almost 
6 million Blueback Herring have been passed through the Holyoke system 
since 1955. Other species that were somewhat unexpected, such as Striped 
Bass (8,400 since 1979) and Gizzard Shad (6,700 since 1986), have been 
lifted in good numbers. The real "king" of the river, the Atlantic Salmon, 
has been slower to reestablish itself, with only 3,000 handled at Holyoke 
since 1977. The numbers of fishes handled at Holyoke between 1969 and 
1996 are shown in Figure 3. Note the general increase in American Shad 
and Blueback Herring through the early 1980s and the lower numbers in 
the mid-1990s. The cause of the decline is poorly understood and may be 
related to a combination of many factors. Some fisheries biologists think it 
might be related to the expanding population of Striped Bass that occurred 
after the mid-1980s (O'Leary 1998, pers. comm.). Of course, once the fishes 
have spawned and the juveniles are free- swimming, they must get back 
downstream. A number of obstacles that are inherently related to dam op- 
eration can kill fishes, including the turbines themselves. Downstream pas- 
sage facilities are already in place at many dams, and their design has been 
the focus of discussion between regulators and dam owners, especially over 
the past five years (M. Tisa 1998, pers. comm.). 

Concurrent with the recent fishway development, there has been an in- 
teragency effort to restore Atlantic Salmon to the Connecticut and Merri- 
mack rivers. Two -year- old juvenile Atlantic Salmon from Canada were 
released in the Connecticut River and the first adult returned in 1974. 
Penobscot River juveniles were released into the river in 1976, and 90 adults 
returned in 1978. As shown in Figure 3, salmon have usually returned in the 
hundreds each year. By the late 1980s, supplemental stocking of salmon fry 



18 Inland Fishes of Massachusetts 




i 1 1 r 



<JD_ SS)> /£> <9 Sn Sn Sg Sg Sg Sg Sn Sg Sg Sg 



1400 
1200 
1000 - 



♦ Atlantic Salmon 
-n — Striped Bass 




Figure 3. Fishes passed at the Holyoke Fish Lift, Connecticut River. 



Ichthyology in Massachusetts 19 



added to the recovery effort, and, by the spring of 1997, the total number of 
fry stocked in the Connecticut Basin was nearly 8.5 million. As mentioned 
in the Atlantic Salmon account, a few instances of natural reproduction in 
the wild have been reported (O'Leary 1997, pers. comm.). 

references. Moffitt et al. 1982, Anon. 1997a, 1997b (Connecticut River 
restoration); Meyer 1999 (recoveries and declines). 



20 Inland Fishes of Massachusetts 



Conserving and Enjoying Fishes 
and the Aquatic Environment 



Freshwater fishes are declining worldwide due principally to degraded or 
changing habitats. The exact reasons for the declines are often difficult to 
attribute to any one cause and usually are tied to multiple factors. The 
aquatic ecosystems of New England have been altered for almost four cen- 
turies; indeed, every body of water in Massachusetts has been negatively 
impacted in one way or another. The physical, chemical, and biological 
characteristics of Massachusetts freshwaters have been changed since the 
colonial days by settlement and subsequent agricultural or industrial ex- 
pansion. Zaitzevsky (1982) notes that, as early as 1645, every marsh within 
the town boundaries of Boston had been modified in some way. Since colo- 
nial times, small- and medium- sized dams, used for grain and sawmills, 
changed the characteristics of the local environment. Virtually all of Massa- 
chusetts was cutover for timber, and 65 percent of the state was fully cleared 
for agriculture by the mid- 1800s. The effects of this massive deforestation 
on the aquatic community due to increased runoff, siltation, and increased 
water temperatures will never be known and can only be presumed. How- 
ever, damming and industrial or urban waste disposal from development 
have had a well-documented effect on local fishes. This is particularly true 
in regard to the large food fishes, such as the Atlantic Salmon, American 
Shad, and sturgeons that return seasonally to the rivers. 

Conservation of North American Freshwater Fishes 

In North America, 3 genera, 27 species, and 13 subspecies of freshwater 
fishes have become extinct over the last century (Williams and Miller 1990). 
At least 300 species, subspecies, or populations are currently listed at some 
level of rarity in North America. Thus, approximately one-third of the North 
American freshwater fish fauna is impacted. Imperilment is not restricted to 
any particular taxonomic group but is higher in areas of greater endemicity 



Conserving and Enjoying Fishes 21 



(Warren and Burr 1994). Even many recognized but not yet scientifically 
described fishes are declining as documented by Williams et al. (1989). 

Massachusetts has only two species on the North American rare fish list 
developed by Williams et al. (1989): the Atlantic and Shortnose sturgeons. 
Both of these species are listed under the Federal Endangered Species Act. 
Seven other species are listed in state categories: the Lake Chub and North- 
ern Redbelly Dace are State Endangered; the American Brook Lamprey and 
a trimorphic freshwater population of Threespine Sticklebacks are State 
Threatened; and the Eastern Silvery Minnow, Longnose Sucker, Bridle 
Shiner, and Burbot are listed as State Special Concern. In addition, a num- 
ber of other species, such as the Common Shiner, have been studied as 
possible list candidates by various private and state agencies. 

It is easier to understand why it is important to conserve the native fresh- 
water fishes of Massachusetts if they are viewed on a national or worldwide 
basis. Most of Massachusetts native fishes belong to a group of fishes found 
along the Atlantic coastal plain from Florida to southern Maine, and many 
of these species are found no place else in the world. The Atlantic coastal 
plain is quite small when viewed globally and also quite endangered; on top 
of it sits the giant East Coast megalopolis. This megacity that once sprawled 
from Boston to Washington, DC, now essentially engulfs the whole span 
from Miami to Portland, Maine, with only remnant natural areas scattered 
along the way. 

Ecological Considerations and Local Fish Distribution 

Fishes are totally integrated with their aquatic world and highly dependant 
on the quality and size of their environment. The distribution of most inland 
fishes is limited by each species' physiological constraints or behavioral re- 
quirements. Some species, such as Slimy Sculpin, Longnose Sucker, and 
salmonids, require relatively cold and clean water. Others, such as White 
Sucker and Brown Bullhead, tend to be more tolerant of a wider range of 
aquatic conditions to live healthy lives. 

Streams and rivers vary naturally according to flow, volume, gradient, sub- 
strate, temperature, and water chemistry. Within these various conditions, 
fishes tend to be distributed in fairly predictable patterns. For example, in 
small, cold Massachusetts headwater streams, Slimy Sculpin and native 
Brook Trout may be the only species found. Farther downstream, Black- 
nose and Longnose dace as well as White Suckers may also be present. Still 

22 Inland Fishes of Massachusetts 



farther downstream, in larger and slightly warmer midreach sections of 
streams, Common Shiners, Fallfish, and Tessellated Darters tend to be- 
come more prevalent, while trout and Slimy Sculpin become scarce. In 
lowland stream environments, warmwater species, such as the Pumpkin- 
seed, Redbreast Sunfish, Brown Bullhead, Golden Shiner, Redfin Pickerel, 
Chain Pickerel, American Eel, Banded Sunfish, and Swamp Darter, become 
more dominant. 

Lakes and ponds throughout the state, as well as impounded sections of 
rivers, also tend to be inhabited by an assemblage of warmwater fish species, 
including Brown Bullhead, Pumpkinseed, Bluegill, Largemouth Bass, Red- 
breast Sunfish, Golden Shiner, Yellow Perch, and Chain Pickerel. Very few 
of the state's lakes and ponds naturally support large numbers of cold- 
water fishes, except the large reservoirs such as Quabbin and Wachusett, 
where introduced Lake Trout and Atlantic Salmon are found. 

The precise combination of species present at any given location depends 
on many factors, but, within any drainage, the number or diversity of native 
fishes from small upland headwaters to larger lowland rivers increases nat- 
urally. However, human activities, including the introduction of non-native 
fish species, pollution, and dams, have had significant impacts on native 
fishes, and, in many drainages, natural patterns of distribution and abun- 
dance no longer exist. Unfortunately, these human-caused disturbances 
are ubiquitous, and even those activities that traditionally have been con- 
sidered "low- intensity" are now known to cause problems. 

A study of a warmwater stream in Virginia (Weaver and Garman 1994), 
for example, has shown that gradual urbanization led to declines in abun- 
dance of all species and trophic groups, and six species were lost during 
the 38-year study period even though no exotic fishes were introduced. 
These low- intensity factors, such as gradual human population increase, 
new homes, new roads, and road crossings at streams, are very likely major 
causes of degraded aquatic ecosystems around all metropolitan areas in 
Massachusetts. 

Other studies of randomly selected northeastern lakes by the U.S. Environ- 
mental Protection Agency (Whittier et al. 1997) showed a regional decline 
in minnow species. This decline was attributed to human development of 
lake shorelines and the presence of non-native predatory fishes. 

The effects of acid precipitation are also a major problem for many of 
Massachusetts' aquatic ecosystems (Halliwell 1985). Above and beyond 
just simply lowering pH, which in itself can cause declines in the survival 

Conserving and Enjoying Fishes 23 



of larvae and eggs and the availability of prey, increased acidification can 
alter water chemistry in numerous ways, including allowing various poten- 
tially toxic metals to more easily enter local ecosystems. A study in eastern 
Massachusetts comparing two ponds, one with low acid-buffering capacity 
and another with higher buffering capacity (Stallsmith et al. 1996), shows 
vulnerability to acid spikes early in a fish's life, as the first gills begin to de- 
velop, along with poor growth rates. 

references. Deacon et al. 1979, Williams et al. 1989 (declining North 
American fishes); Miller et al. 1989 (extinct North American fishes); Wil- 
liams and Miller 1990 (conservation status); Foster 1992, 1999 (land use, 
MA); Haines and Baker 1986 (acidification). 

Fishwatching and Fish Photography 

Humans have always been fascinated with the behavior of fishes and have 
woven them into mythology and folklore. When one thinks of watching 
fishes, the first thing that comes to mind is the coral reef environment; 
however, ponds and streams in Massachusetts offer good opportunities to 
observe and photograph a number of interesting species. 

Observing Fishes Fishwatching is by no means a new idea, but today's 
cameras, underwater housings, strobe lights, wet suits, face masks, snorkels, 
scuba, and miniature tape recorders have opened worlds below water that 
were impossible to access 50 years ago. Even with today's equipment, 
fishwatching starts with patience and the ability to sit quietly Many fishes 
are easy to see; nesting sunfishes can be watched by standing on the shore 
at almost any pond during early summer. Anadromous fishes, such as the 
Alewife, can be seen in incredible numbers as they pass up fishways on 
both small and large streams. The general public is allowed to view the 
major fishways on the Connecticut and Merrimack rivers. Other fishes are 
more difficult to see, and the watcher may have to sit still on a bank for half 
an hour or more or watch from a distance with binoculars. But even the 
wary species will return if they become gradually acclimated to the pres- 
ence of the motionless observer. As many anglers will attest, you can often 
see small minnows and darters by walking slowly in streams. 

For the more adventurous, a face mask and a snorkel can be used to study 
fishes in almost every aquatic habitat in Massachusetts. In many areas it is 



24 Inland Fishes of Massachusetts 



not necessary to get into deep water. Small hill streams may be cold but of- 
fer views of sculpins, trout, and dace. Despite the fact that warmwater ponds 
and streams are often murky, the world of the Yellow Perch, Pumpkinseed, 
and Largemouth Bass can be viewed. If you are really lucky, Banded Sun- 
fish, Swamp Darters, and Redfin Pickerel can be seen between the weeds. 
Safety should always come first. Cold water can lower body temperatures, 
and many local rivers, streams, lakes, reservoirs, and ponds are littered with 
dangerous objects ranging from glass to automobiles. 

For the less adventurous, it is possible to keep and observe many of our 
local native fishes in home aquaria. Warmwater species such as catfishes, 
sunfishes, and killifishes do well indoors and are interesting to watch. Cau- 
tion should be exercised in selecting what fishes to keep. It is best to have 
some experience with fishes before bringing any home from the wild be- 
cause many fishes will feed only on live food and are difficult to keep alive. 
Local fish and game laws must also be observed, and, in some cases, per- 
mits may be necessary. Contact a Massachusetts Division of Fisheries and 
Wildlife office to obtain the current regulations. 

Photographing Fishes Fishes can be photographed either in the wild by 
swimming with underwater cameras or by placing a fish in an aquarium at 
streamside. A valid state fishing license or a special permit is required to 
catch and hold any wild fish. Fishes cannot be moved from the collecting 
site or released elsewhere without additional permits. Aquaria with gravel 
bottoms and other natural features can often produce excellent photo- 
graphs. Strobes should be used to stop action, but all light sources must 
be set at an angle to the aquaria to prevent reflections. 

Technical photographs used to illustrate morphological characteristics 
generally show a preserved specimen in a uniform, left-sided view without 
habitats in the background. The fish is placed between a glass plate and the 
front of an aquarium and photographed. Photographing the fishes in water 
eliminates surface reflections and allows a record of detailed scales and fin- 
rays. Live fishes can be photographed in a similar manner, but care should 
be taken not to stress the fish. Vegetation and backgrounds can be added to 
create natural effects. 

With the advent of several models of underwater cameras, it is now pos- 
sible to get into the water and quietly approach your subject. With the 
proper equipment to keep warm and the patience to move slowly, you can 



Conserving and Enjoying Fishes 25 



take good photographs. It is best to swim or crawl upstream so that any silt 
suspended by your movements will wash away. 

references. Emery and Winterbottom 1980, Holm 1989, Jenkins and 
Burkhead 1993 (photography); McDonald et al. 1972 (watching fishes) ; 
Mills 1990 (keeping and photographing fishes); Quinn 1990 (keeping native 
fishes). 



26 Inland Fishes of Massachusetts 



The Land, Water, and Fishes 
of Massachusetts 



The Land 

The political region we call Massachusetts sits atop an area of New England 
that is a transition zone between the warm coastal plain to the south and 
the boreal forest to the north. Due to its geographical position, Massachu- 
setts' physical and natural features are diverse, and its flora and fauna are 
elements of the ocean, the coastal plain, and the mountains. A wonderful 
overview of the state's natural biomes is presented in The Nature of Massa- 
chusetts (Leahy et al. 1996), which includes chapters on aquatic communities 
such as salt marshes, coastal plain ponds, freshwater marshes, lakes and 
ponds, rivers and streams, and the floodplain forest, all important to fishes. 
Located in the northeastern portion of the United States, Massachusetts 
covers 8,257 square miles. It is the third most densely populated state in the 
United States but ranks only forty- sixth in total area. According to the 1993 
US census, the 1990 population was over 6 million with an average density 
of 730 people per square mile. Elevations across the state range from 3,491 
feet at Mount Greylock in northwestern Berkshire County to sea level along 
the shore. The coastal plain of eastern Massachusetts has elevations that 
are generally less than 250 feet. 

The Water 

The Major River Drainages The watersheds of Massachusetts can be di- 
vided into 9 major basins or drainage areas and 33 smaller drainages (see 
Figure 4, Map 2, Table 2; Halliwell et al. 1982). An overview of these water- 
sheds, their flora and fauna, and other characteristics can be found in An 
Atlas of Massachusetts River Systems (Bickford and Dymon 1990). Most of 
the major river drainages in Massachusetts flow from or into other New En- 
gland states. For example, New England's largest river basin, the Connecti- 
cut, originates at the Canadian border and flows south, separating Vermont 



The Land, Water, and Fishes of Massachusetts 27 




Figure 4. The major drainage basins of New England. 



28 Inland Fishes of Massachusetts 




Map 2 Massachusetts drainages: 11 Hoosic; 12 Kin- r$< 

derhook; 13 Bashbish; 21 Housatonic; 31 Farmington; 

32 Westfield; 33 Deerfield; 34 Connecticut; 35 Millers; 

36 Chicopee; 41 Quinebaug; 42 French; 51 Blackstone; 

52 Ten Mile; 53 Narragansett; 61 Mount Hope; 62 Taunton; 

71 Mystic; 72 Charles; 73 Neponset; 74 Weymouth- Weir; 

81 Nashua; 82 Concord; 83 Shawsheen; 84 Merrimack; 91 Parker; 

92 Ipswich; 93 North Shore; 94 South Shore; 95 Buzzards Bay; 96 Cape 

Cod; 97a Martha's Vineyard; 97b Nantucket (after Halliwell et al. 1982). 



.,- 







"96 






from New Hampshire, then moves through Massachusetts and Connecticut, 
and finally empties into Long Island Sound. In total, there are more than 
2,027 streams and rivers that traverse approximately 5,465 miles (8,795 km) 
in Massachusetts. The Connecticut and Merrimack river basins are the larg- 
est, draining 2,726 and 1,200 square miles respectively. The smaller coastal 
drainages form a large combined watershed that drains approximately 
2,352 square miles. 

The major Massachusetts watersheds are outlined below and on Map 2. 
The area of each drainage, the number of ponds, and the surface acreage of 
the ponds are shown in Table 2. 

Hudson River Basin contains the Hoosic, Kinderhook, and Bashbish 
drainages that flow north and west out of Massachusetts into the Hudson 
River and then south to the Atlantic Ocean. 

Housatonic River Basin originates in western Massachusetts and flows 
south through Connecticut into Long Island Sound. 

Connecticut River Basin includes the Farmington, Westfield, Deerfield, 
Millers, and Chicopee drainages in Massachusetts. The basin drains south 
from the Canadian border to Long Island Sound. 

Thames River Basin includes the Quinebaug and French drainages in Mas- 
sachusetts, which flow south through Connecticut to Long Island Sound. 

Merrimack River Basin flows south from New Hampshire and contains 



The Land, Water, and Fishes of Massachusetts 29 



Table 2 Massachusetts water resource summary. 



Basin 



Drainage 



Map Code* Sq. Miles No. Ponds Acres 



HUDSON 


Hoosic 


11 


166 


17 


625 




Kinderhook 


12 


22 


4 


28 




Bashbish 


13 


16 


5 


41 


HOUSATONIC 


Housatonic 


21 


500 


118 


5184 


CONNECTICUT 


Farmington 


31 


156 


50 


3583 




Westfield 


32 


517 


79 


4550 




Deerfield 


33 


347 


21 


589 




Connecticut 


34 


669 


103 


2992 




Millers 


35 


313 


104 


4720 




Chicopee 


36 


723 


166 


31837 


THAMES 


Quinebaug 


41 


154 


55 


2406 




French 


42 


95 


66 


3603 


MERRIMACK 


Nashua 


81 


443 


161 


10953 




Concord 


82 


400 


127 


7456 




Shawsheen 


83 


78 


19 


552 




Merrimack 


84 


279 


79 


4033 


MASSACHUSETTS 


Parker 


91 


82 


17 


316 


BAY 


Ipswich 


92 


155 


74 


1982 




North Shore 


93 


172 


90 


2428 




Mystic 


71 


76 


45 


1496 




Charles 


72 


319 


142 


3708 




Neponset 


73 


117 


61 


1879 




Weymouth-Weir 


74 


91 


27 


1213 




South Shore 


94 


241 


173 


5151 


SOUTHERN NEW 


Blackstone 


51 


335 


185 


7267 


ENGLAND 


Ten Mile 


52 


49 


42 


1301 




Narragansett 


53 


56 


5 


146 




Mount Hope 


61 


56 


7 


3713 




Taunton 


62 


530 


213 


12551 




Buzzards Bay 


95 


375 


171 


6378 




Cape Cod 


96 


403 


343 


11039 




Islands 


97 


160 


102 


7453 



* Refers to Map 2 

three drainages in Massachusetts. The Nashua Drainage courses north 
from Massachusetts through New Hampshire, and the Concord Drainage 
(the combined Concord- Sudbury- Assabet system) and Shawsheen Drain- 
age flow into the Merrimack main stem before it enters the Atlantic Ocean 
at the northeast corner of Massachusetts. 



30 Inland Fishes of Massachusetts 



Massachusetts Bay Drainage Area contains a large assemblage of drain- 
ages flowing to the Massachusetts Bay portion of the Gulf of Maine, north 
of Cape Cod. Included are the Parker, Ipswich, North Shore, Mystic, Charles, 
Weymouth-Weir, Neponset, and South Shore drainages. 

Southern New England Drainage Area is a group of drainages that, with 
the exception of a few streams on Cape Cod, drain south into Narragansett 
or Buzzards Bay and Nantucket Sound. Included are the Blackstone, Ten 
Mile, and Narragansett drainages, all with headwaters in Massachusetts, 
that flow south through Rhode Island to Narragansett Bay; the Taunton and 
Mount Hope drainages, completely in Massachusetts, which drain south 
into Mount Hope Bay; and the Buzzards Bay, Cape Cod, and Island drain- 
ages, which include Martha's Vineyard and Nantucket. 

The Fishes 

Fossil Fishes Fossil fish records from inland New England date back almost 
200 million years. Remains of the extinct genera Semionotus, Redfleldius, 
and Diplurus (see Figure 5) can be found in sedimentary rocks from huge 
lakes in what is now the Connecticut River Valley. The genus Semionotus 
is the most commonly found local fish fossil; based on the frequency with 
which its fossils are seen, this genus existed in large numbers. Redfleldius, 
a more elongate fish, is much less commonly encountered. However, the 
rarest of all, at least in the Connecticut Valley, is Diplurus, a member of 
the lobe-limbed fish group that some scientists think might have given rise 
to tetrapods. One species of the lobe-finned fishes, called the Coelocanth, 
Latimeria, is still living today in the deep marine waters of the Indian Ocean. 

More recent fossils are absent, primarily because of the nature of New 
England's geological events. However, based on archaeological informa- 
tion, the Massachusetts fish fauna of 3,000 to 700 years ago probably con- 
tained the same fish species as when the Pilgrims arrived. Scarce records 
from archaeological sites, usually fragments of scales, vertebrae, otoliths, or, 
rarely, pharyngeal and buccal jaw elements, indicate that the Native Ameri- 
cans ate sturgeons, river herrings, Chain Pickerel, Fallfish, and Brook Trout. 

references. Colbert 1970; Olsen 1980; Olsen and McCune 1991. 

Recent Fishes Currently 83 species of fishes, both native and introduced, 
from 27 families have been documented as regular residents of Massachu- 



The Land, Water, and Fishes of Massachusetts 3 1 




Figure 5. The Connecticut River Valley fossil fishes. Top: Semionotus (re- 
drawn from Olsen and McCune 1991). Middle: Redfieldius (redrawn from 
Schaeffer and McDonald 1978). Bottom: Diplurus (redrawn from Schaeffer 
1952). 



setts freshwaters (see Table 3 and Appendix 2). This book outlines all spe- 
cies that permanently live in freshwater and also those that usually move 
between fresh- and saltwater during various stages of their life histories. It 
does not cover the occasional vagrant marine species that sometimes enter 
the edge of freshwater. 

The species treated in this book can be separated into three ecological 
groups: 1) primary freshwater fishes that live their entire life cycles in fresh- 
water; 2) secondary freshwater fishes that have the physiological capacity 



32 Inland Fishes of Massachusetts 



Table 3 Number of reproducing species per drainage basin. 

Basin Primary Secondary Introduced Diadromous Total 



Hudson 


15 





8 





23 


Housatonic 


22 





16 





38 


Connecticut 


26 


1 


26 


7 


60 


Thames 


17 





14 





31 


Merrimack 


21 


8 


18 


10 


57 


Mass. Bay 


18 


12 


12 


9 


51 


S. New England 


19 


18 


12 


10 


59 


Islands 


11 


17 


2 


4 


34 


State Total 


29 


18 


27 


10 


84 



to move back and forth between salt- and freshwater; and 3) diadromous 
fishes that make relatively long seasonal migrations between salt- and 
freshwater. Most diadromous fishes are anadromous; like the herring and 
salmon, they are born in freshwater, grow at sea, and migrate to freshwater 
as adults to spawn. The American Eel is catadromous, with a life cycle the 
reverse of the salmon and herring. Eels are born at sea, return to freshwater 
to grow, and then re-enter the sea to spawn and die. Table 3 shows the rela- 
tive numbers of species from each group found in each drainage. 

Two native fishes have been extirpated from the Massachusetts portions 
of their range. First, the Atlantic Salmon, although currently being reintro- 
duced, disappeared in the mid- 1800s after the construction of dams. The 
second species, the Trout-perch, was known from the Hoosic and Housa- 
tonic drainages and was last found in the early 1940s. 

Surprisingly, 48 percent of Massachusetts primary fish species are not 
native to the state but were introduced and are now reproducing in local 
waters. These species are mostly game fishes and include black basses and 
sunfishes, pike, and several catfishes, which were introduced to enhance 
sportfishing. Other non-native species include some minnows that escaped 
or were released from baitbuckets. In fact, the introduced Bluntnose Min- 
now, unknown from Quabbin before 1984, was the most abundant shore- 
fish in the reservoir during our autumn surveys in 1989. The 27 exotic spe- 
cies now reproducing in Massachusetts are treated in detail in the species 
accounts. 

In addition, a fair number of nonreproducing exotic species have been 
reported from local waters (Hartel 1992, Cardoza et al. 1993). Reports of 

The Land, Water, and Fishes of Massachusetts 33 



non-native North American species include a few observations of a true 
gar, probably the Spotted Gar, Lepisosteus oculatus, one record of a North- 
ern Hog Sucker, Hypentelium nigricans, and three species of minnows, 
including Emerald Shiner, Notropis atherinoides, Grass Carp, Ctenopharyn- 
godon idella, Red Shiner, Cyprinella lutrensis (Hartel 1992), and Mosquito - 
fish, Gambusia affinis. The following fishes from outside North America 
have all been found in Massachusetts: the pacu-like Pirapatinga, Piractus 
brachypomus, and tambaqui, Colossoma macropomum, which are both 
from South America; the Grass Carp, Ctenopharyngodon idella, and Walk- 
ing Catfish, Clarias batrachus, from Asia; the Oscar, Astronotus ocellatus, 
from South America; the Midas Cichlid, Cichlasoma citrinellum, from Cen- 
tral America; the Giant Snakehead, Channa micropeltes, from Asia; and one 
of the African Upside-down Catfishes, Synodontis. Most surprising, more 
than a dozen documented records (and other verbal reports) of the Red Pi- 
ranha, Pygocentrus natter eri, have been brought to our attention over the 
last decade. Almost all of these fish were caught by anglers and brought to 
the Massachusetts Division of Fisheries and Wildlife or the Museum of 
Comparative Zoology for identification. Fortunately, most of these exotic 
fishes cannot survive winter water temperatures. However, a second spe- 
cies of Asian shakehead, Channa argus, was collected live from Newton 
Pond, Shrewsbury, in 2001. This species ranges quite far north in Asia and 
could survive Massachusetts winters. 

Non-native or exotic fishes, those introduced from outside their native 
drainages, from both near and far, have been involved in the decline of 
many native species worldwide (Courtenay and Stauffer 1984). The exotic 
fishes found in Massachusetts are mostly released aquarium pets, but 
some, like the Grass Carp, were the result of deliberate, though illegal, 
introductions. 



Origin and Distribution of the Fauna 

The origin of native fishes found today in Massachusetts can be traced back 
to a period just after the last glaciers receded from the Northeast around 
14,000 years ago. These large bodies of ice covered the Northeast for about 
70,000 years and made the survival of freshwater vertebrates impossible. All 
traces of the fish species that lived here shortly before the glaciers have been 
lost to time. However, by understanding the geological events and the geo- 
graphical ranges of native North American species, we can postulate how 



34 Inland Fishes of Massachusetts 




Figure 6. The glaciated Northeast about 14,000 YBP. Dotted line shows the present 
coastline; a solid line indicates the coastline at that time; and a solid line with perpen- 
dicular bars shows the extent of glaciation (adapted from Emery 1987). 



freshwater fishes moved into New England as the glaciers melted and re- 
treated to the north. 

As the glacial ice retreated, Cape Cod and the islands of Martha's Vine- 
yard and Nantucket were formed from the material that accumulated at the 
edges of the glaciers. The sea level dropped almost 100 feet during maximum 
glaciation, and large areas of exposed continental shelf formed a broad 
coastal plain along eastern North America (see Figure 6) that contained 
abundant streams created by glacial melt. Mastodons and mammoths 
roamed this coastal plain, part of which is now Georges Bank. 

Ancestors of the native species found in Massachusetts today survived 
the glacial period in areas south of the ice. These areas, called "refugia," 



The Land, Water, and Fishes of Massachusetts 35 




Figure 7. The receding glacier about 11,500 YBP. Proto- Georges Bank is an island; ice 
blocks are present at what will be Long Island Sound and near Stellwagen Bank 
(adapted from Schmidt 1986, Emery 1987). 



allowed the genetic stocks to survive and spread into areas once covered by 
ice. The routes that the fishes employed in populating Massachusetts pose 
intriguing questions. 

The Southern Invasion The current distribution pattern of fishes along the 
Atlantic coastal plain indicates that the majority of Massachusetts fresh- 
water fish species survived the glacier in refugia along the coast, possibly as 
far south as North Carolina (Schmidt 1986), or in areas off the present Con- 
necticut coast (Whitworth 1996). A northern coastal refugium in the vicinity 
of Georges Bank has also been suggested, and this area might account for 
some of the forms found on Cape Cod and the Islands. At various times, 
Georges Bank was a cape (14,000 YBP) or an island (11,500 YBP; see Figure 7) 
that may have harbored isolated populations of plants, invertebrates, or ver- 
tebrates in southern New England (D.G. Smith 1992, pers. comm.; Schmidt 
1986: 148). Variation in some northern forms of estuarine fishes, such as 
Mummichogs, Rainwater Killifish, and Inland Silversides, may be related to 
this phenomenon. 
It is postulated that, as temperatures moderated, fishes from the refugia 



36 Inland Fishes of Massachusetts 



gradually migrated into New England rivers and streams. They invaded in- 
land to all areas except where physical or ecological barriers, such as water- 
falls, stopped their passage. Some species, such as the Swamp Darter and 
Banded Sunfish, were possibly prevented from entering some of the south- 
ern New England drainages by a giant residual ice block thought to be pres- 
ent in what is now Long Island Sound (Schmidt 1986) or by the rise of salt 
water into glacial Lake Connecticut (Whitworth 1996). 

The Northern Route The distribution of a few northern New England 
fishes seems to be most closely linked to the Great Lakes fauna, which sur- 
vived glaciation in the Mississippi Valley. Although there is little geological 
evidence to support the theory or indicate the actual route followed, it is 
likely that species, such as the Redbelly Dace, followed glacial streams, bogs, 
and flooded lakes that probably linked the precursors of the upper Hudson 
and Saint Lawrence basins to today's Connecticut River Drainage and other 
areas of northern New England. Once into the upper headwaters of the 
Connecticut, their route into Massachusetts would have been easily accom- 
plished. Species such as the Lake Chub, Burbot, and Trout-perch may also 
have arrived from the north. 

Current Distribution The recent distribution of fishes in Massachusetts is 
shown on the maps found adjacent to each species account and summa- 
rized in Appendix 2. The numbers of native primary species historically 
known from the larger Massachusetts drainage basins range from 21 to 26 
(see Table 3), whereas the medium- sized drainage areas have only 15 to 19. 
The islands of Martha's Vineyard and Nantucket have far fewer native pri- 
mary species, and it is difficult to prove which of those on the islands are 
native or transplanted from the mainland. The numbers of species per 
drainage area and the species composition are, in part, related to historic 
geology in that some species just were not able to migrate into drainages 
having appropriate habitats. However, certain regions, like parts of the 
Massachusetts Bay Drainage Area, have certainly lost species due to factors 
related to urbanization. No clear-cut zoogeographic pattern emerges from 
this simple analysis except that a few coastal plain species, such as the 
American Brook Lamprey, Redfin Pickerel, and Banded Sunfish, are not 
found in the western part of the state. Conversely, the Redbelly Dace, Lake 
Chub, Eastern Silvery Minnow, Creek Chub, Longnose Sucker, and Trout- 
perch are found only in the western areas. Others like the Slimy Sculpin, 

The Land, Water, and Fishes of Massachusetts 37 



Blacknose Dace, Longnose Dace, and Fallfish are common to the west but 
show strong indications that they were more widely spread to the east in re- 
cent historic times. 

The only clear differentiation is in the number and distribution of native 
secondary species, with more species found in the Southern New England 
Drainage Area. There are about twice as many native secondary species 
south of Cape Cod, especially if the Bay Anchovy and Inland Silverside, 
which are rare north of the Cape, are considered part of the southern fauna. 
This faunal break follows a well-documented distributional pattern for ma- 
rine fishes where many mid-Atlantic species are not found north of Cape 
Cod or Georges Bank. 

Explanations of how or why species are found in some drainages and not 
in others are difficult. The problem lies in interpreting current distribution 
data that have gaps and misleading information due, at least in part, to hu- 
man involvement during the last four centuries. Eighteenth-century extir- 
pation, transplantation, or movement through built canals from drainage 
to drainage confuse the data. In addition, the fact that comprehensive sur- 
veys of the state's freshwater fauna were not made until the 1940s does not 
allow the establishment of baseline data on which to evaluate patterns of 
distribution. 

references. Raymo and Raymo 1989 (New England geology); Schmidt 
1986, Whitworth 1996 (New England fish zoogeography); Emery 1987 
(Georges Bank). . 



38 Inland Fishes of Massachusetts 



How to Use This Book 



Identifying Fishes 

As in animal tracking or birdwatching, looking at fishes requires the devel- 
opment of good powers of observation. However, since there are relatively 
few species of fishes in Massachusetts, identification is somewhat sim- 
plified. One method is to look at illustrations or color plates and match 
them with a fish. Careful attention should be paid to the number, size, and 
placement of the fins as well as to the size of the scales, the general body 
shape, and color pattern. The various features of a fish are diagrammed in 
Figure 8. A review of the distribution maps and the descriptions of habitats 
should also help in identification by eliminating some species. After match- 
ing a fish to an illustration, the next important step is to read the identifica- 
tion section as it outlines each species' salient features. 

The best way to identify a fish is to use the identification keys. The family 
key (page 51) will identify a fish to its family and will direct the reader to a 
species key. If only one member of a family occurs in Massachusetts, the 
family key will identify it directly to the species. 

Identification keys in this book are presented in pairs of illustrated state- 
ments called "couplets." Each part of a couplet gives one or several choices, 
which are opposite those given in the other part of the couplet. For example, 
the "a" part of the couplet might state that the fish has an adipose fin, while 
the "b" part of the same couplet states that the fish lacks such a fin. The 
reader chooses the one that best describes the fish to be identified. The key 
then leads to another couplet and ultimately to the correct identification. 
Careful reading of the couplets is essential. 

Special equipment is not necessary, although a hand lens or an inexpen- 
sive microscope may be needed to identify smaller fishes, particularly min- 
nows. In addition, a small probe or a large needle will help to count or sep- 
arate features such as fin rays, gill rakers, or scales. 



How to Use This Book 39 



Uniform and accurate counts and measurements are important in identi- 
fying fishes. The standard methods of measuring fishes described by Hubbs 
and Lagler (1964) and Jenkins and Burkhead (1993) are followed in this book. 
We have kept terminology and the types of counts and measurements to a 
minimum in the keys and identification section. A section with selected 
counts containing key information is given at the beginning of each species 
account (the counts are principally from reviews of Trautman 1981, Scott 
and Crossman 1973, Smith 1985, and Jenkins and Burkhead 1993). For a full 
set of counts and measurements, C.L. Smith's The Inland Fishes of New 
York State (1985) covers most of the Massachusetts species. Under selected 
counts, the following abbreviations are used: D = dorsal fin, A=anal fin, 
GR=gill rakers, Pec=pectoral fin, Pel=pelvic fin, PT=pharyngeal teeth, 
Vert = vertebrae. Scales are usually indicated by three sets of numbers sepa- 
rated by forward slashes. They indicate scales above the lateral line /scales 
along the lateral line /scales below the lateral line. Uppercase Roman nu- 
merals indicate true spines, lowercase Roman numerals indicate spinelike 
soft rays, and Arabic numerals indicate true soft rays. 

General Anatomy 

The overall anatomy of the body parts of a fish is presented in Figure 8, with 
two fishes to illustrate most of the body parts mentioned in this book. In 
addition, the head is shown in detail to demonstrate its parts and the gill 
arch structure, which can be seen by lifting the gill cover. 

Measurements 

As shown in Figure 8, there are two common ways of measuring fishes. 
The most common is total length (TL), often used by anglers and fisheries 
biologists. Total length is the maximum length of the fish from the furthest 
projections of the jaws or snout to the tip of the tail. The other type, called 
standard length (SL), is used for more precise measurements. With this sys- 
tem, the fish is measured from the tip of the upper jaw to the base of the 
bony plate that supports the tail fin. This point on the tail often shows a 
crease in the skin when the tail is bent. Standard length allows the accu- 
rate measurement of fishes even when the tail is damaged. Head length 
(HL) is measured from the tip of the upper jaw T to the posterior edge of the 
gill cover. 



40 Inland Fishes of Massachusetts 



Counts 

Rays and Fin Spines The elements supporting the fins are generally of two 
types, either soft rays or hard spines (see Figure 8), which can be counted 
easily by placing a light behind the fin. 

Soft rays are usually, but not always, branched and flexible. Most rays are 
bilaterally paired and segmented. Occasionally, soft rays are hard struc- 
tures, as in the spines of carp or catfishes, and are often called spines but 
are actually hardened soft rays. They can always be identified as rays if 
they are divided, branched, or segmented. Soft rays can usually be easily 
counted by following these rules: In minnows (Cyprinidae), suckers 
(Catostomidae), and trout (Salmonidae), only the principal rays are 
counted. These include the single, large, unbranched ray at the front of 
the fin and the remaining branched rays. In catfishes (Ictaluridae) and a 
few other groups, with fins that taper forward and result in small anterior 
rudimentary rays, all rudimentary rays are counted; and, in catfishes, the 
skin along the base of the anal fin often has to be removed to expose the 
small rays. Always count the last two rays of the dorsal fins as one when 
they are joined together at their bases. 

Fin spines are unsegmented and usually hard and sharp but may be flex- 
ible as in the sculpins (Cottidae). All spines are counted, even the smallest. 

Scales Scale counts are often the only way to separate closely related spe- 
cies, and accurate counts can usually be made only on preserved material. 
For small fishes, a microscope may be needed, and it is best to let the scales 
dry slightly to see the scale edges. Another method is to direct a small jet of 
air at each scale as it is counted, using a hollow needle on a rubber hose 
connected to a small aquarium pump. 

Lateral-line scale counts are made along the lateral line or along the mid- 
body of a fish that does not have a lateral line. The lateral line is a sensory 
system, and scales along it have a pore or tube that leads to a sensory de- 
vice. Counts start at the posterior end of the opercle and end at the base of 
the caudal fin. The base of the caudal fin is the area where a crease forms 
when the tail is folded back and forth. 

Scales above the lateral line are counted diagonally, downward, and back- 
ward, in a row from the dorsal fin origin to the lateral line. The count follows 
the natural row of scales and includes the small scales at the base of the 
dorsal fin, but not the lateral-line scale. 



How to Use This Book 41 



Barbels: 

1. nasal 

2. chin 

3. maxillary 



dorsal 
spine 



dorsal fin 



adipose fin 




caudal fin 



anal fin 



pelvic fin 



TOTAL LENGTH 




scales 
above 
lateral line 



caudal 
peduncle 



pectoral fin 

pelvic fin 



anal fin 



scales 
below 
lateral line 



premaxilla 



mandible 




opercular 
spine 




maxilla opercle 



subopercle 



gill filaments 



Figure 8. General anatomy of a fish. 



42 Inland Fishes of Massachusetts 



Scales below the lateral line are counted upward and forward from the 
beginning of the anal fin. 

Scales before the dorsal fin are counted along the midline of the back 
from the origin of the dorsal fin to the area at the rear of the head where the 
scales stop. All scales that intercept the midline are counted. 

Gills Gill rakers, the structures on the anterior surface of the gill arch, 
should not be confused with the gill filaments, which are used for respira- 
tion and are found on the posterior gill arch (see Figure 8). The gills can be 
observed by lifting the opercular flap, which may have to be cut open along 
the bottom in some fishes (especially catfishes). Gill raker counts usually 
include all of the rakers, even the smallest most rudimentary ones, on the 
first gill arch. However, in some cases, the keys will indicate that a count of 
only the rakers on the lower half of the arch is required. 

Interpreting the Distribution Data 

Information about the distribution of Massachusetts fishes is the key part 
of this book. These data, represented on the maps, are based on examina- 
tion of museum specimens, field surveys, and literature. The original data 
for these maps are available for examination at the Museum of Compara- 
tive Zoology, Harvard University, or at the Field Headquarters of the Mas- 
sachusetts Division of Fisheries and Wildlife, Westborough. 

Museum Specimens Well over 50,000 museum specimens were examined 
to verify fish identifications and distribution. A few at the Museum of Com- 
parative Zoology date back to the mid- 1800s, but most were collected after 
1950. Museum specimens form the most solid database, especially for the 
smaller, more difficult to identify species. Specimens used in this study are 
stored principally at the Museum of Comparative Zoology, Harvard Univer- 
sity (MCZ), and the Museum of Zoology, University of Massachusetts, Am- 
herst (UMA). Additional material at Northeastern University, Boston; South- 
eastern Massachusetts University, Dartmouth; the University of Michigan, 
Ann Arbor; Fitchburg State College; and Cornell University, Ithaca, was also 
examined. Map 3 shows the areas from which the museum specimens were 
collected. 

The MCZ has some older specimens collected by D.H. Storer, W.H. Put- 
nam, R.H. Wheatland, and S.F. Baird from the mid- 1800s. A small series of 



How to Use This Book 43 




Map 3 Museum specimens: localities from which 
museum specimens have been examined. 



fv^v^W^K - r «? * 






specimens from B. McCabe's 1942 thesis on the 
stream fishes of western Massachusetts was transferred 
from Springfield College to the MCZ in 1979. More recent "* 

accessions include R.H. Gibbs' Boston University collections from the 
early- 1960s and a large amount of material collected by K.E. H artel, princi- 
pally with C.R. Gougeon, D.G. Smith, TJ. Andrews, and A.E. Launer, between 
1975 and 1989. A large number of voucher specimens from the statewide 
surveys by D.B. Halliwell and other Massachusetts Division of Fisheries and 
Wildlife (MDFW) workers (1977-1990) have also been deposited at the MCZ. 

The collections at the University of Massachusetts, Amherst, were made 
almost single-handedly by Prof. T.J. Andrews from 1948 to 1980, and some 
newer material has been acquired by D.G. Smith, since 1974. The University 
of Massachusetts museum contains some of the best reference material 
from western Massachusetts. 



2f\ 



•■ 



Fisheries Surveys In addition to museum specimens, MDFW field surveys 
have helped document distribution. The early surveys were designed pri- 
marily to sample game fishes, but, in some cases, information on selected 
nongame fishes was also included; later survey work routinely listed all fish 
species encountered. 

Stream and River Surveys Britton McCabe's work (1942, 1943, Map 1) on 
the stream fishes of the Hudson-Hoosic, Housatonic, and Connecticut 
drainages began a series of stream surveys that include the Westfield River 
(Mullan 1952); the Millers and Squannacook (Mullan 1953); the Merrimack- 
Ipswich (Schlotterbeck 1954); and the Taunton-North (Bridges 1955). A 
summary of these surveys was completed by Tompkins and Mugford (1964). 



44 Inland Fishes of Massachusetts 
















•■-•-• 



NW 



Map 4 Stream surveys: localities of stream surveys V C*/ t£/"\ y ^ \J I cJ^s P" 
by Massachusetts Division of Fisheries and Wildlife, x Jj§ 7 

1969-1990 (see Halliwell 1989). 



> \ 



eg v s 



2U< 



Since 1969, additional surveys have been con- ^ ,£ s;i 

ducted by the state with federal aid funds. These more w* ... 

recent surveys (Map 4) include the Charles (Bergin 1969); Housatonic 
(Bergin 1970); Chicopee (Bergin 1972); Blackstone (Bergin 1973a); Deerfield 
Green (Bergin 1973b); Nashua (Madore 1974); Taunton (Madore 1975); 
Hoosic and Farmington (Madore 1976); and Westfield (Halliwell 1978). A 
statewide stream survey and classification project was initiated by the 
MDFWin 1979. During this study, records from 1,430 collections made at 
691 streams statewide were analyzed and form the basis of a Ph.D. disserta- 
tion by D. Halliwell (1989). 






Lake, Pond, and Reservoir Surveys The first set of lake and pond surveys 
was conducted by the MDFW from 1942 to 1952 and covered 385 lakes 
statewide (Swartz 1944; McCabe 1948, 1952, 1953; McCabe and Swartz 1952; 
Stroud 1955; Map 5). Along with the surveys from 1952 to 1968, 76 lakes 
across the state were chemically treated to remove unwanted fish species 
and facilitate trout management. Additional MDFW surveys covering an- 
other 385 lakes (436 samples) were carried out between 1960 and the early 
1990s, but these studies were not published. However, selected information 
from these surveys is available in pond booklets printed by the Massachu- 
setts Division of Fisheries and Wildlife. These booklets include pond maps 
and are available at minimal cost from the division's Westborough Field 
Headquarters. Data based on both the published pond data and the un- 
published field data were compiled, reviewed, and used for this book. 



How to Use This Book 45 




fcati 






Map 5 Lake and pond surveys: localities of lakes -{(/ i \/ ~^J% ijl \J% 

and ponds surveyed by Massachusetts Division of 
Fisheries and Wildlife; open circles 1960-1976; closed 
circles 1977-1989. 



! JJV (\ s 

y \/ J \\ >■ 



■jlrfc 



9 o, 



), > £•', 






. y * 



A full list of all the ponds in Massachusetts can be found 
in Ackerman et al. (1984). 

Between 1991 and 1996, an additional 27 Massachusetts lakes were 
monitored by the US-EPA Environmental Monitoring and Assessment 
Program (Whittier et al. 1997). 






' ^?"\ 



Coastal Rivers and Estuaries Data for distribution of upper estuarine spe- 
cies have been obtained from a review of the bay and estuary surveys of the 
Massachusetts Division of Marine Fisheries that were published in their 
Monograph Series between 1965 and 1975. Clayton et al. (1978) reviewed 
data for many coastal species using both published and unpublished data, 
including a summary of these surveys. 

Distribution Maps Each symbol on a map represents a locality from 
which we have seen specimens or carefully reviewed reports and literature 
accounts. A single symbol may represent several closely situated sites. Most 
records are of museum specimens or from MDFW field surveys; in most 
cases, marine records are not shown. On some maps, two types of symbols 
are shown to give information, as explained in the caption under the map. 
Solid or open circles are most often used to show the presence or absence 
of a species in recent surveys. In maps that present this information, the 
solid symbols represent all verified records; however, many of these sites 
were sampled only once, possibly many years ago. Open circles show sites 
where a species was documented before 1970 but where it was not found 
when resampled during recent studies (1975-1991). Several open circles, 



46 Inland Fishes of Massachusetts 



especially if clumped together, may indicate loss of a species from that area 
or an environmental problem. However, these data should be interpreted 
cautiously because of possible sampling errors. 

OVERVIEW AND OUTLINE OF FAMILY 
AND SPECIES ACCOUNTS 

Family Accounts 

Each family account outlines a group of closely related species. Scientific 
names of families always end in the suffix "idae." The sunfish family, Cen- 
trarchidae, the pike family, Esocidae, and the salmon family, Salmonidae, 
are three examples. The family accounts provide general information on 
such topics as worldwide distribution, relationships to similar fishes, and 
the number of species in the family. Technical terms are defined in the glos- 
sary (Appendix 3). 

Species Accounts 

These sections give the common and scientific names of the species, their 
conservation status, a line drawing, and a reference to the appropriate 
color plate. Also included are sections on identification, size, natural his- 
tory, distribution and abundance, special notes, and references. 

Names. Names used in this book generally follow those suggested by the 
American Fisheries Society's Common and Scientific Names of Fishes from 
the United States and Canada (Robins et al. 1991a), except when changes 
have been published since that release. The common name of each species 
appears on the first line of each species account. Below it is the scientific 
name, consisting of two words, first the generic and then the specific name. 
These names are always italicized in print. Following the species name is 
the name of the person who described the species, along with the date of 
the description. When the describer's name and date appear in parenthe- 
ses, it indicates that the species is now placed in a different genus from the 
original description. Scientific names for the fish species treated in this 
book are not given in the text because they are noted at the beginning of 
each species account. However, species that are not treated in this book 
have the scientific name used each time they are mentioned. 



How to Use This Book 47 



Species Status. At the beginning of each species account the status of each 
species is given. These categories (native or introduced) are based on docu- 
mented introductions, literature reviews, and our interpretation of each 
species' distribution in North America. Also, the status of rare species (spe- 
cial concern, threatened, or endangered) is noted at the head of each spe- 
cies account. 

Size. Information on total length (TL), measured in inches, is given in all 
accounts. In some accounts, we supplement TL with a metric standard 
length (SL) to document or clarify records. 

Distribution and Abundance. This section describes the state and local 
range, as well as general abundance, and includes a map for each species. 
Information on the total range of these species is available in Lee et al. (1980 
et seq.) and in Page and Burr (1991). For an explanation of distribution data 
and symbols on maps, see page 43. 



48 Inland Fishes of Massachusetts 



Identification Keys and Species Accounts 



Key to the Families and Monotypic Species 
of Massachusetts Inland Fishes 



la. Jaws absent (mouth is an "oral disk"); 
pectoral fins absent; seven gill openings 
on each side of head. Lamprey family, 
Petromyzontidae, page 60. 



lb. Jaws present (may be modified into 
"sucker mouth"); pectoral fins present; 
one gill opening on each side of head. 
Go to 2. 




2a. Caudal fin strongly asymmetrical; 
several rows of bony scutes along body. 
Sturgeon family, Acipenseridae, page 66. 

2b. Caudal fin roughly symmetrical, 
lower lobe less pronounced; if bony 
scutes present, they are found in one row 
along midbody line. Go to 3. 




3a. Both eyes on same side of head; body 
extremely flattened. American Sole fam- 
ily, Achiridae. One local species: Hog- 
choker, Trinectes maculatus, page 280, 
Plate 55. 




*yyf*y*;'-r-y-i>T' 



3b. Eyes on opposite sides of head; body 
not extremely flattened. Go to 4. 




Family and Species Accounts 5 1 



4a. Pelvic fins absent; body very elon- 
gate. Go to 27. 




4b. Pelvic fins present; body sometimes 
slightly elongate. Go to 5. 




5a. Fleshy adipose fin present. Go to 15. 




5b. Adipose fin absent. Go to 6. 




6a. Single median barbel on underside of 
lower jaw. Cod family, Gadidae, page 189. 



6b. Barbels, if present, are paired on 
either side of jaws. Go to 7. 




52 Inland Fishes of Massachusetts 



7a. Single dorsal fin with either no spines 
or with one serrated "spine" at anterior 
edge; no pelvic spines. Go to 18. 




7b. Either one dorsal fin with distinct 
"spine" and "ray" portions or two dis- 
tinct fins (first may be series of "free 
spines"); pelvic fin with one or more 
spines. Go to 8. 




8a. First dorsal fin consisting of series of 
"free spines." Stickleback family, Gas- 
terosteidae, page 219. 



8b. Spines in first dorsal fin connected 
by membranes. Go to 9. 




9a. First and second dorsal fins sepa- 
rated by distance equal to or greater than 
the length of the base of the first dorsal 
fin; origin of pelvic fins well behind base 
of pectoral fins. Go to 14. 



9b. First and second dorsal fins sepa- 
rated by distance less than the length of 
the base of the first dorsal fin, or are 
united; origin of pelvic fins directly below 
or in front of the base of the pectoral fins. 
Go to 10. 





,.-.---^)))))))))));; ))>)) ; )))))))))))) - 





Family and Species Accounts 53 



10a. Anal spines two or fewer (spines 
may be flexible) . Go to 1 1 . 

10b. Anal spines three or more. Go to 12. 




10b 



1 la. All anal fin elements flexible, no 
stout fin spines. Sculpin family, Cottidae. 
One local freshwater species: Slimy 
Sculpin, Cottus cognatus, page 232, 
Plate 54. 



lib. Anal fin with one or two inflexible 
spines. Perch and darter family, Percidae, 
page 266. 





12a. Deeply forked caudal fin with nar- 
row peduncle; bony scutes present on 
caudal peduncle. Jack family, Carangi- 
dae. One local species in freshwater: 
Crevalle Jack, Caranx hippos, page 277. 



12b. Caudal fin shallowly forked with 
relatively broad peduncle; no bony 
scutes on caudal peduncle. Go to 13. 




54 Inland Fishes of Massachusetts 



13a. Preoperculum has serrated poste- 
rior edge with spine; first and second 
dorsal fins separated; anal fin squared. 
Striped Bass family, Moronidae, page 235. 



13b. Preoperculum has smooth poste- 
rior margin without spine; first and sec- 
ond dorsal fins united, at least by short 
membrane; anal fins rounded. Sunfish 
and Black Bass family, Centrarchidae, 
page 241. 




14a. Mouth upturned; origin of second 
dorsal fin well behind origin of anal fin. 
Silverside family, Atherinopsidae, page 
211. 

14b. Mouth horizontal; origin of second 
dorsal fin directly over origin of anal fin. 
Mullet family, Mugilidae, page 216. 




15a. Barbels present; scales absent. Bull- 
head Catfish family, Ictaluridae, page 143. 



15b. Barbels absent; scales present. Go 
to 16. 




Family and Species Accounts 55 



16a. Dorsal fin with two spines; mouth 
does not extend to front margin of eye; 
scales with rough posterior edge. Trout- 
perch family, Percopsidae. One local spe- 
cies: Trout-perch, Percopsis omiscomay- 
cus, page 186, Plate 30. 



16b. Dorsal fin with no spines; mouth 
extending beyond front margin of eye; 
scales with smooth posterior edge (cy- 
cloid). Go to 17. 




17a. Pelvic axillary process present; 
more than 100 scales in lateral line series. 
Trout family, Salmonidae, page 171. 




17b. Pelvic axillary process absent; fewer 
than 80 scales in lateral line series. Smelt 
family, Osmeridae. One local species: 
Rainbow Smelt, Osmerus mordax, page 
168, Plate 43. 




18a. Caudal fin rounded or squared. 
Go to 25. 



18b. Caudal fin at least partially forked. 
Go to 19. 



56 Inland Fishes of Massachusetts 



19a. Origin of dorsal fin almost directly 
above origin of anal fin. Go to 24. 




19b. Origin of dorsal fin distinctly in 
front of origin of anal fin. Go to 20. 




20a. Scales along midline of belly form- 
ing a sharp edge or "keel." Herring fam- 
ily, Clupeidae, page 78. 




20b. Midline of belly without "keel." 
Go to 21. 




21a. Maxilla extends well past posterior 
margin of eye. Anchovy family, Engrauli- 
dae. One local freshwater species: Bay 
Anchovy, Anchoa mitchilli, page 89. 




21b. Maxilla does not extend past 
middle of eye. Go to 22. 




Family and Species Accounts 57 



22a. Serrated "spine" at anterior edge of 
dorsal fin. Minnow family (in part), 
Cyprinidae, page 92. 



22b. All dorsal fin elements flexible. 
Go to 23. 




23a. Dorsal fin with nine or fewer rays; 
lips smooth; pharyngeal teeth in one or 
two rows with nine or fewer teeth per 
row. Minnow family (in part), 
Cyprinidae, page 92. 



23b. Dorsal fin with 10 or more rays; lips 
with "pleats"; pharyngeal teeth in one 
row with over 20 comblike teeth. Sucker 
family, Catostomidae, page 135. 




24a. Snout broad and flattened; needle- 
like beak absent; pectoral fins ventral; 
dorsal fin rounded. Pickerel family, Eso- 
cidae, page 157. 

24b. Head narrow and round in cross 
section; needlelike beak present; pectoral 
fins lateral; dorsal fin falcate. Needlefish 
family, Belonidae. One local freshwater 
species: Atlantic Needlefish, Strongylura 
marina, page 195. 




58 Inland Fishes of Massachusetts 



25a. Dorsal fin long (more than 45 rays); 
gular plate present. Bowfin family, Ami- 
idae. One species: Bowfin, Amia calva, 
page 71. 




25b. Dorsal fin short (fewer than 15 
rays); gular plate absent. Go to 26. 




26a. Upper jaw not protrusible; groove 
between premaxillaries and snout not 
continuous. Mudminnow family, Umbri- 
dae. One local species: Central Mudmin- 
now, Umbra limi, page 165, Plate 29. 



26b. Upper jaw protrusible; groove be- 
tween premaxillaries and snout continu- 
ous. Killifishlike families, Cyprinodonti- 
dae and Fundulidae, page 198. 





27a. Body covered with bony rings; 
mouth modified into tubelike structure; 
dorsal, caudal, and anal fins distinct. Sea- 
horse and pipefish family, Syngnathidae. 
One local species: Northern Pipefish, 
Syngnathus fuscus, page 229. 

27b. Body not covered with bony rings; 
mouth not tubelike; dorsal, caudal, and 
anal fins continuous. Freshwater eel 
family, Anguillidae. One local species: 
American Eel, Anguilla rostrata, page 75. 




&** 





Family and Species Accounts 59 



Lamprey Family 

Petromyzontidae 



Lampreys and the marine hagfishes are the only surviving jawless fishes. 
Lampreys have a number of unusual morphological features, including an 
oral disk that is surrounded by a fleshy hood in the larval stage, horny (kera- 
tinized) teeth in the adult, seven pairs of gill openings, and a single nostril 
(nasohypophseal opening) on the dorsal midline in front of the eyes. In ad- 
dition, they lack pectoral fins, pelvic fins, scales, true jaws, and ossified 
skeletons. Lampreys include 41 species in six genera. 

The life history of all lampreys is divided into two ecologically and mor- 
phologically distinct parts: the larva, often termed an ammocoete, and the 
adult. The ammocoete lives in freshwater, typically in sheltered backwaters 
of rivers and streams that have sand and detritus substrates. They are rela- 
tively small, blind, filter-feeding burrowers. Depending on the species, adult 
lampreys are either parasitic and often live for several years, or nonparasitic 
(actually nonfeeding as adults) and live only a short period after reproduc- 
ing. Adult lampreys construct shallow, round nests by picking up and ar- 
ranging stones on the streambottom with their oral sucking disks. Sea Lam- 
preys supported a substantial fishery during Colonial days in New England. 

references. Hardisty 1979 (biology); Hardisty and Potter 1971-82 (taxon- 
omy, distribution, biology). 



Key to Massachusetts Lampreys 



la. Teeth absent, eyes covered with skin, 
oral hood present. Larval stage, ammo- 
coetes, go to 2. 




lb. Teeth present, eyes developed, 
mouth rounded and exposed. Adult 
stage, go to 3. 




& & & G> 0© 



60 Inland Fishes of Massachusetts 



2. Key to Larvae 

2a. Unpigmented postnostril area ap- 
proximately equal to the size of nostril; 
most of lip pigmented; area above 7 gill 
openings mostly pigmented. Sea Lam- 
prey, Petromyzon marinus, page 64. 




2b. Unpigmented postnostril area about 
twice the size of nostril; most of lip un- 
pigmented; wide unpigmented band 
above gill openings. American Brook 
Lamprey, Lampetra appendix, page 62. 




3. Key to Adults 

3a. Mouth with many teeth in multiple 
rows; mature adults generally over 2 feet 
in length. Sea Lamprey, Petromyzon mar- 
inus, page 64, Plate 2. 




3b. Mouth with scattered horny teeth, 
multiple rows only in anterior portion (if 
at all); adults less than 1 foot in total 
length. American Brook Lamprey, Lam- 
petra appendix, page 62, Plate 1. 




Family and Species Accounts 61 



American Brook Lamprey 

Lampetra appendix (DeKay 1842) 



Native, State Threatened 

PLATE 1 




identification. Lampreys have seven pairs of gills but lack true jaws, 
pectoral fins, and pelvic fins. Adult American Brook Lampreys seldom grow 
as large as 8 inches total length (TL), while mature Sea Lampreys are usu- 
ally over 24 inches TL. Adults of the two species also have distinctive ar- 
rangements of the horny teeth in the oral disk (see key Figure 3). The larvae 
(ammocoetes) can be separated by examining the pigment pattern: Ameri- 
can Brook Lampreys lack areas of pigment on the side of the head, on the 
lips of the oral hood, around the nostril, and along the side of the body 
above the gill openings (see key Figure 2). 

size . Larvae of American Brook Lampreys generally transform to adults at 
4 to 6 inches TL. The largest specimen reported from Massachusetts is a 
transforming individual, 6.75 inches TL. 

natural history. American Brook Lampreys begin to transform into 
the nonparasitic adult form in the late summer, maturing by late winter or 
early spring. Soon after completing metamorphosis, the adults construct 
shallow nests in sandy gravel and spawn from mid-April to early May in 
Massachusetts. The eggs hatch in nine days at 68°F and, after several days, 
the young (ammocoetes) burrow into areas of soft substrate. They remain 
as larvae for four to five years and filter feed mainly on small algae and a 
variety of other microorganisms found in detritus. Adults do not feed and 
die shortly after spawning. 

distribution and abundance. In Massachusetts, this species is 
known only from the eastern Blackstone River Drainage, the Mashpee River 



62 Inland Fishes of Massachusetts 




American Brook Lamprey. 



* 



&'»» 



on Cape Cod, and the Mill and Tiasquam rivers 
on Martha's Vineyard. American Brook Lampreys 
are not common. In fact, the first Massachusetts 
specimens, found in the Blackstone Drainage in early 
1950, were misidentified as Sea Lampreys. Finally, in the mid-1960s, a spec- 
imen was collected and correctly identified by J. Musick (Virginia Institute 
of Marine Science) and J. Hoff (South Eastern Massachusetts University) 
while they were trout fishing in the Mashpee River. The Martha's Vineyard 
population, which appears stable, went unnoticed until our surveys in 1988. 



c 



p 



'/'■' 



notes. The American Brook Lamprey is listed as a threatened species in 
Massachusetts because of its limited distribution and the species' sensitiv- 
ity to environmental change. This species requires streams with clean, silt- 
free water, riffle areas for nesting, and backwaters with detritus beds for 
larval growth. 

references. Halliwell 1979 (Massachusetts); Hoff 1988 (Mashpee River); 
Moore and Beamish 1973 (habits); Rohde et al. 1976 (life history, Delaware); 
Vladykov and Follet 1967; Vladykov and Kott 1980 (descriptions, key); 
Vladykov 1973 (conservation). 



Family and Species Accounts 63 



Sea Lamprey Native 

Petromyzon marinus Linnaeus 1758 plate 2 




identification. Lampreys have seven pairs of gills but lack true jaws, 
pectoral fins, and pelvic fins. Sea Lampreys are similar to but much larger 
than American Brook Lampreys; Sea Lampreys are usually over 24 inches 
TL as adults. Adults of the two species also have distinctive arrangements of 
the horny teeth in the oral disk (see key Figure 3) . The larvae (ammocoetes) 
can be separated by examining the pigment pattern; Sea Lampreys have 
much more pigment on the side of the head, on the lips of the oral hood, 
around the nostril, and along the side of the body above the gill openings 
(see key Figure 2). Male Sea Lampreys, in breeding condition, develop a 
prominent ridge along the dorsal midline and a cloacal appendage. These 
features are absent in the females. 

size. Ammocoetes of Sea Lampreys begin to transform into juveniles at 4 
to 8 inches TL. Adults, returning from the sea, are from 24 to 34 inches TL at 
the Holyoke Dam on the Connecticut River. 

natural history. Adult Sea Lampreys return to freshwater after spend- 
ing at least two years feeding at sea. While in the ocean, they are parasitic 
and attach themselves to a variety of fishes with their oral disk and feed al- 
most exclusively on the body fluids of the host. Sea Lampreys migrate up- 
river to small- and medium-sized streams with gravel and rocky substrates 
in late May or early June as water temperatures reach 50° to 59°F. They re- 
main near the breeding sites for several weeks before spawning but do not 
feed. Sea Lampreys construct shallow nests by picking up small stones with 
their oral disks. During spawning, the female attaches to a rock on the up- 
stream edge of the nest and releases between 124,000 and 305,000 eggs over 
several days. The adult Sea Lampreys die shortly after spawning. The eggs 
hatch in approximately two weeks, when the young ammocoetes drift 

64 Inland Fishes of Massachusetts 








Sea Lamprey: coastal records are not indicated. 

downstream to burrow into areas of soft substrate. 

They remain in freshwater for four to five years and M- 

filter feed on a wide variety of organisms and detritus. & "'" /^^ ° I 

distribution and abundance. Prior to the 1800s, Sea Lampreys en- 
tered virtually every Massachusetts stream and river that allowed them ac- 
cess to breeding sites. In the mid- 1800s, newly constructed dams blocked 
their migration routes and industrial pollution altered their habitat. Re- 
cently, new fishways constructed for anadromous fish runs have allowed 
them to return to many areas. Sea Lampreys are now common in the Con- 
necticut River (up to 53,000 per year at the Holyoke Fish Lift) and migrate 
north of the Massachusetts border. The first recent records from the Millers 
Drainage date from the late 1980s, just after the opening of the Turners Falls 
Fishway. Sea Lampreys are also locally common in portions of the Merri- 
mack and Parker rivers. They are much less frequently encountered in the 
South Shore, Cape Cod, and other coastal drainages. 

notes . Although Sea Lampreys have caused significant damage to the 
fisheries in several large lakes (notably the Great Lakes), they have no nega- 
tive effect on the inland fisheries of Massachusetts. The adults and juve- 
niles generally do not feed while in freshwater. However, if Sea Lampreys 
were to invade any of Massachusetts' larger reservoirs, such as Quabbin, 
the effects might easily mirror the negative impacts that have occurred in 
the Great Lakes. Ammocoetes are an important part of stream food webs 
because they feed on detritus. 

references. Beamish 1980 (biology); Bigelowand Schroeder 1948, Miller 
1980 (distribution, natural history); Stier and Kynard 1986a (abundance, 
size, sex ratio, MA), 1986b (spawning, MA). 



Family and Species Accounts 65 



Sturgeon Family 

Acipenseridae 



Sturgeons are part of an ancient group of fishes that is at least 200 million 
years old and is closely related to the curious-looking paddlefishes of the 
Mississippi Drainage and China. Sturgeons are characterized by five rows 
of bony shields or scutes along the body, a cartilaginous internal skeleton, 
an intestine with a spiral valve, a heterocercal tail, and a ventral protrusile 
mouth preceded by four barbels. These fishes are typically anadromous, al- 
though some of the world's 25 species live their entire lives in freshwater. 
Sturgeons are generally sluggish but strong fishes that can leap 4 to 6 feet 
out of the water. Unfortunately, many sturgeon species are declining. All of 
the North American species are listed as endangered, threatened, or of spe- 
cial concern in various parts of their ranges. 

references. Binkowski and Doroshov 1985 (biology and management); 
Lauder and Liem 1983 (relationships); Vladykov and Greeley 1963 (review); 
Deacon et al. 1979, Ono et al. 1983 (threatened species); Kologe 1992 (MA); 
Birstein et al. 1997 (biology, relationships, conservation). 



Key to Massachusetts Sturgeons 



la. No bony plates between base of anal 
fin and lateral row of scutes; mouth over 
60 percent of interorbital width; intestine 
and peritoneum dark. Shortnose Stur- 
geon, Acipenser brevirostrum, page 67, 
Plate 3. 




lb. A row of 2 to 6 small bony plates 
between base of anal fin and lateral row 
of scutes; mouth less than 50 percent 
interorbital width; intestine and peri- 
toneum light in color. Atlantic Sturgeon, 
Acipenser oxyrinchus, page 69, Plate 3. 




66 Inland Fishes of Massachusetts 



Shortnose Sturgeon 

Acipenser brevirostrum Lesueur 1818 



Native, Federally Endangered 
plate 3 




identification. Shortnose Sturgeon lack small bony plates, or scutes, 
between the base of the anal fin and the lowest row of primary scutes, and 
the viscera and lining of the body cavity are blackish. The snout is short and 
blunt in adults, but its length varies with age and may be relatively long and 
pointed in juveniles. Young Shortnose and Atlantic sturgeon can be distin- 
guished by the relative width of the mouth (see key Figure la). Shortnose 
Sturgeon never grow larger than 4.5 feet, compared with the 12 to 14 feet 
attained by Atlantic Sturgeon. 

selected counts. D 38-42; A 19-22; GR 13-25. 

size. The largest Massachusetts specimens are about 40 inches TL and 
weigh 20 pounds. 

natural history. Shortnose Sturgeon are often anadromous, but the 
Massachusetts populations appear to be mostly riverine. In general, Short- 
nose Sturgeon move upriver in the fall, then overwinter, and spawn in early 
May. After the spawning season, individuals move downriver, but the 
movements are complicated because some fishes in each population do 
not spawn each year. Adults do not spawn until they are almost 10 years 
old. The oldest documented Massachusetts specimen was 37 years old. 
Shortnose Sturgeon are bottom feeders with a variable diet; sturgeons of all 
sizes eat crustaceans and insects, whereas larger individuals eat hard- 
shelled invertebrates, such as mussels and snails. 

distribution and abundance. In Massachusetts, the most stable 
populations are in the Connecticut River, where two populations exist with 
little interchange. One is landlocked between the Turners Falls Dam and 



Family and Species Accounts 67 







Shortnose Sturgeon: open circles indicate historic x \y I '%/""- .5 ^ ul ijl )- 
records where the species may no longer occur; oce- 
anic records are not indicated. 




m'AXrx .1, 



i w 



s / -Aa 



\£" 



the Holyoke Dam and the other ranges downstream £ \f\ 

,._,/ct%..«'.^ ~> 

of the Holyoke Dam. The total number of adults in the ° ,-? 

Connecticut River is thought to be fewer than 1,000 fish. The first confirmed 
Merrimack record of this species is based on a juvenile specimen collected 
in 1956 that we found in the Cornell University collection. Shortnose Stur- 
geon were studied in detail in the lower Merrimack by the Massachusetts 
Cooperative Fishery Research Unit in 1988. Older records indicate that 
populations existed in Waquoit Bay, Cape Cod, the Taunton and Parker 
rivers, and possibly the Charles River. 



V, 

A 



notes. The Shortnose Sturgeon is rare throughout its range and is listed 
as endangered by the United States Fish and Wildlife Service. Historically, 
damming, overfishing, and pollution have contributed to the decline of the 
species. 

references. Buckley and Kynard 1981 (spawning), 1985a (movement), 
1985b (habitat and behavior); Dadswell et al. 1984, Kynard 1997 (review); 
Gorham and McAllister 1974 (identification); Kieffer and Kynard 1993, 1996 
(Merrimack); Ono et al. 1983 (threatened species); Tracy 1906 (Taunton 
River); Taubert 1980 (reproduction). 



68 Inland Fishes of Massachusetts 



Atlantic Sturgeon Native, State Endangered 

Acipenser oxyrinch izs Mitchill 1814 plate 3 




identification. Atlantic Sturgeon are distinguished from Shortnose 
Sturgeon by a row of small scutes above the anal fin and by their light- col- 
ored viscera. The two species can also be separated by the relative width of 
the mouth (see key Figure lb). The snout tends to be long, narrow, and 
slightly upturned, but its length varies with age. 

selected counts. D 38-46; A 26-28; GR 16-27. 

size. This is Massachusetts' largest freshwater fish. Historic records from 
1634 mention sturgeon of 12, 14, and 18 feet. The 18-foot record is probably 
an exaggeration, but sturgeons might have grown that large in the past. 
Typically, Atlantic Sturgeon reach 6 to 9 feet. 

natural history. Historically abundant, this species declined before 
the turn of the century and almost nothing is known about its natural his- 
tory in Massachusetts. In other areas, Atlantic Sturgeon are anadromous, 
with adults moving upriver to spawn over clay, rubble, gravel, or shell bot- 
toms in brackish to freshwater in mid-May to mid-June. The smallest 
spawning females are close to 20 years old, 6.5 feet long, and 1 10 pounds. 
The juveniles usually remain in freshwater for three to four years, or until 
they reach about 30 inches TL. Older juveniles and adults leave the estuar- 
ies in the fall and migrate south along the coast. Young Atlantic Sturgeon 
feed principally on soft benthic invertebrates, whereas the adults feed on 
larger invertebrates, such as mollusks. In saltwater, they will eat crabs, 
worms, and large numbers of Sand Lance Ammodytes. 

distribution and abundance. In Massachusetts, the species is rare, 
and recent records are limited to the Merrimack and Taunton rivers. In the 
mid- to late 1970s, a single, 8-foot-long sturgeon was seen at the base of the 

Family and Species Accounts 69 




■JA±^~ \ \ /^.i..' a - W ...../ r —\ :_/: L2sUxm xi \ i.\. -e^ j\ ,\\\j \T-\ <r*? 



Atlantic Sturgeon: open circles indicate historic 
records where the species may no longer occur; 
oceanic records are not indicated. 




f # 4, 



Essex Dam on two occasions during late June. Sur- 
veys between 1988 and 1991 by the Massachusetts Co- 
operative Fisheries Unit have studied and radio-tagged several 30-inch fish 
in the vicinity of Haverhill. Whether these fish are part of a reproducing 
Merrimack population or are just transients is unknown. Historic records 
indicate that they reproduced in the Taunton River; three juveniles were 
found there by the Massachusetts Cooperative Fisheries Research Unit in 
1991. Specimens are still found in small numbers along the coast and are 
occasionally taken by trawlers. There are also old records that indicate that 
this species entered the Charles, the Parker, and other coastal rivers. 






notes. Historically, this species migrated up the Merrimack River as far 
as Amoskeag Falls in Manchester, New Hampshire, but the dams erected 
in the mid- 1800s prevented this annual movement. Probably the last large 
catch of Atlantic Sturgeon was in 1887, when two tons were taken on the 
Merrimack River in one week. 

references. Bigelow and Schroeder 1953 (marine records); Murawski 
and Pacheco 1977, Hoff 1980 (synopsis); Jerome et al. 1965 (Merrimack 
population); Tracy 1906, Buerkett and Kynard 1993 (Taunton River); Kieffer 
and Kynard 1993 (Merrimack River). 



70 Inland Fishes of Massachusetts 



Bowfin Family 

Amiidae 



The Amiidae is an ancient family of ray-finned bony fishes that was much 
more diverse 140 to 200 million years ago. Although they have an extensive 
fossil record, there is only one living species, the Bowfin of North America. 
They have a long dorsal fin, a gular plate between the lower jaws, hetero- 
cercal tail, and small tubelike anterior nostrils. In addition, Bowfins possess 
a lung-like gas bladder that enables them to breathe air and occupy habi- 
tats that lack the dissolved oxygen required by most other fishes. They also 
have numerous stout teeth on their jaws and gill arches. As "living fossils," 
Bowfins are popular in sciences classes, both as an animal for experiments 
and for anatomical dissections. Bowfins have been of great importance in 
the analysis of the evolution of vertebrates and the subject of numerous 
anatomical and behavioral investigations. 

references. Boreske 1974 (fossil history); Lauder and Liem 1983; Grande 
and Bemis 1998 (relationships). 



Bowfin Introduced 

Amia calva Linnaeus 1766 







identification. The Bowfin is easily recognized by its long body, long 
dorsal fin with more than 45 rays, and gular plate. Small Bowfins might be 
confused with Central Mudminnows since they superficially resemble each 
other, but the latter lacks a gular plate and has a much shorter dorsal fin. 
A dark spot on the upper base of the tail is prominent in juvenile and male 
Bowfins. 



Family and Species Accounts 71 




■^.-i...L..L«c.\... 



Bowfln. 

selected counts. D 42-58; A 9-10; 
Scales 64-68. 



size. Bowfins are quite large; individuals over 30 inches 
long are common in some parts of their range. 



natural history. Bowfins are typically found in vegetated backwaters 
of rivers but are also found in cool, clear bodies of water. Bowfins are often 
nocturnal or crepuscular. The males clean a nest area 1.5 to 3 feet in diame- 
ter in water less than 3 feet deep and spawn in the spring. Nests are some- 
times quite close, and males vigorously defend the nest area and its eggs or 
young. Newly hatched young adhere to the surrounding vegetation with an 
adhesive organ on the top of their heads. After young Bowfins leave the 
nest, they form schools that are continuously protected by adult males for 
several weeks. Bowfins are voracious feeders; fishes are their primary food 
source, but they will eat almost any animal they encounter, including 
crayfishes and frogs. 

distribution and abundance. Endemic to North America, this spe- 
cies is found west and south of the Hudson River. The source of the intro- 
duction to Massachusetts waters is unknown. Prior to the late 1980s, the 
only record from Massachusetts was of a single Bowfin specimen taken 
from Lake Onota in Pittsfield, during 1974. In 1986, however, a number of 
specimens were found in an impoundment in Easthampton, and the fol- 
lowing year a large specimen was taken from the Connecticut River in Sun- 
derland. A specimen was also reported from the fishlift at Lawrence on the 
Merrimack River in 1986. An adult male and female were identified by R. 
Hartley (MDFW) from Lake Waldo, Brockton, in 1996. In 1992, a 4-inch ju- 



72 Inland Fishes of Massachusetts 



venile was dropped by a Belted Kingfisher along the Connecticut River in 
Hadley in 1992, which confirmed reproduction in the wild. 

notes. Bowfins are just one of many types of fishes that have evolved vari- 
ous methods of breathing air. Bowfins use their lung-like gas bladder, as do 
lungfishes. Other groups of fishes respire using modifications of their buc- 
cal cavity, and some swallow air to extract the oxygen in the intestines. 

references. Lagler and Hubbs 1940 (diet); Lauder 1980 (feeding); 
MacKay 1963 (reproduction); Reighard 1900 (natural history) , 1902 (repro- 
duction). 



Family and Species Accounts 73 



Freshwater Eel Family 

Anguillidae 



Eels belong to the order Anguilliformes, which contains some 25 families 
and more than 600 species. Eels are elongate, snakelike fishes without pel- 
vic fins or fin spines. The tarpons and bonefishes do not look like eels but 
are members of this same order. Both true eels and tarponlike fishes have 
a specialized larva called a "leptocephalus" that links the two groups. The 
freshwater eels, Anguillidae, are probably the most familiar and commer- 
cially important. They are one of the most generalized eel groups and have 
small imbedded scales that are lacking in almost all other eels. Freshwater 
eels are catadromous; that is, the eggs hatch in the sea, the young migrate 
to freshwater to grow, and the adults return to the sea to spawn. Both the 
American and European eels breed southwest of Bermuda; then the larvae 
make their way back to the coasts of America and Europe. The adults have 
never been captured or seen in the breeding area, and the exact depth and 
method of spawning remains a biological mystery. 

Freshwater eels are considered a delicacy in Europe and Japan. They are 
commercially important in Europe, where about 15,000 metric tons are 
harvested each year. These eels are not used as much in North America; the 
annual harvest in the United States is only 2,000 metric tons, most of which 
is exported to Europe. 

references. Schmidt 1922 (discovery of the breeding area); Tesch 1977 
(general); Ege 1939, Smith 1989 (review and description). 



74 Inland Fishes of Massachusetts 



American Eel Native 

Anguilla rostrata (Lesueur 1817) 




identification. Eels can be identified by their elongate, snakelike bod- 
ies, single small gill openings, true jaws, and pectoral fins. The dorsal fin 
begins far behind the pectorals in American Eels, which distinguishes them 
from the Conger Eel, Conger oceanicus, which is found in Massachusetts 
marine waters. Color varies in eels: at sea, larval eels are nearly transparent 
and colorless and, as they first assume adult shape, retain their transparency 
and are called "glass eels." Upon reaching freshwater, the larvae gradually 
develop pigment to become bronze-black above and silver-white below as 
adults. 

selected counts. D 240; A 200; Vert 103-111. 

size. Female American Eels may grow to over 4 feet TL and weigh up to 
16.5 pounds. A 52-inch female, weighing 7 pounds 8 ounces, and with a 
girth of 7.5 inches, was taken on hook and line from Santuit Pond, Mash- 
pee. Males are much smaller than females, usually 12 to 14 inches TL. Any 
American Eel over 16 inches TL is undoubtedly a female. 

natural history. American Eels that live in Massachusetts are spawned 
in the open ocean south of Bermuda. After hatching, the larval eels begin a 
yearlong journey to New England; presumably, they are carried northward 
on the ocean currents. Off the continental shelf, they begin transformation 
into a 2- to 2.5-inch, transparent, adultlike glass eel. Beginning in March, 
glass eels enter the estuaries and assume adult coloration. Many of these 
juveniles, or elvers, remain in the estuaries, but many thousands migrate 
hundreds of miles up rivers. Only the largest dams stop them, since they 
can usually crawl up cracks and crevices in the face of small dams. Eels live 



Family and Species Accounts 75 




1 /...iLucJ 



American Eel: coastal records are not indicated. 



IS 



«1 






in fresh or brackish water for 7 to 20 years; inver- f •) 

tebrates, fishes, and carrion make up a large part * 

of their diet. When mature, the eels stop feeding and 

in the autumn begin a nocturnal movement to the sea. As 

they migrate, many body changes take place; most noticeably, they become 

bronze-black above and silver-white below and the males' eyes almost 

triple in size. 

Reproductively mature eels have never been seen in the wild, but labora- 
tory studies show that even more body changes take place. Teeth are lost, 
the pectoral fin becomes longer, the gas-producing properties of the swim- 
bladder and associated capillaries change, and even the retinal pigments 
adapt to the low blue light of the ocean. The fully ripe females are almost 
nothing more than a bag of eggs. A large female may contain up to several 
million eggs in a belly so swollen that it appears that the eel can hardly 
function. Almost surely, eels die shortly after spawning, but adults have 
never been seen in the spawning area. The eels' tolerance of habitats that 
range from freshwater streams to open ocean over a mile deep, as well as 
their odyssey as larvae and again as adults, make this drab-looking fish 
truly remarkable. 



4\ 



?>•-'*. 






distribution and abundance . American Eels are common along 
the Massachusetts coast as well as in ponds, rivers, and streams that are 
connected to the ocean. Eels travel overland at night in wet weather and 
can move long distances underground in pipes and culverts, which is why 
they are found in ponds or lakes that appear to lack connections with the 
sea. Though American Eels are still common, a rangewide study by Alex 
Haro (Conti Anadromous Fish Research Center) and colleagues shows 
declines in populations between 1984 and 1995. 



76 Inland Fishes of Massachusetts 



notes. Eels were a very valuable commercial Massachusetts fishery. Over 
240,000 pounds were taken in 1919; however, by 1947, only 33,000 pounds 
were taken annually. Catches have varied through the years since then due 
to economy and demand rather than the actual abundance of eels. In the 
late 1970s, a large export fishery existed but this market collapsed with the 
changing dollar and with the discovery of pollutants in eel flesh. Currently, 
only a small local market for eels as food and bait exists in Massachusetts. 

references. Bigelow and Schroeder 1953 (life history); Haro and Krueger 
1988 (pigmentation, size, migration); Smith 1989 (systematics, life history, 
physiology). Haro et al. 2000 (population declines). 



Family and Species Accounts 77 



Herring Family 

Clupeidae 



The herring family, closely related to the anchovies, contains about 
180 species in 56 genera. The family is united by having a well-developed 
lateral-line system on the head that extends onto the operculum and a spe- 
cialized stethoscopelike connection between the gas bladder and the skull. 
Many herrings have laterally compressed bodies, with thin, loosely at- 
tached scales, and are usually silver. Their teeth are small or absent, and 
their gill rakers are often numerous, long, and thin. While often found in 
marine environments, many herrings enter rivers to spawn, and some spe- 
cies live completely in freshwater. Herrings have a worldwide distribution 
and are extremely abundant in many coastal regions. They are primarily 
planktivores, but some species will eat larger prey, including small fish. 
Small prey are usually filtered out of the water, but larger prey are individu- 
ally picked out of the water column. The family includes all of the fishes 
commonly marketed as sardines. Clupeids are one of the world's most 
commercially fished groups. In the early 1980s, they accounted for approxi- 
mately 30 percent of the world's marine fish harvest, with 14 million tons 
taken each year. 

The Hickory Shad, Alosa mediocris, is included in the identification key 
but not in the species accounts because it does not enter freshwater in 
Massachusetts. Other herrings, including the Atlantic Herring, Clupea 
harengus, and the Atlantic Menhaden, Brevoortia tyrannus, are common 
in Massachusetts coastal waters but seldom stray near the lower edge of 
freshwater. 

references. Bigelow and Schroeder 1953 (general, Gulf of Maine); Hilde- 
brand 1963 (Western Atlantic) ; Whitehead 1985 (review, general biology, 
andsystematics). 



78 Inland Fishes of Massachusetts 



Key to Massachusetts Herrings and Shad 



la. Mouth terminal, last dorsal ray not 
elongate, adults with fewer than 75 lower 
gill rakers. Go to 2. 



lb. Mouth slightly subterminal, last dor- 
sal ray elongate, adults with more than 
100 lower gill rakers. Gizzard Shad, Doro- 
soma cepedianum, page 86, Plate 7. 




2a. Only 18 to 24 gill rakers on lower limb 
of first gill arch; lower jaw strongly pro- 
jecting. Hickory Shad, Alosa mediocris. 
See family account. 



2b. More than 35 gill rakers on lower 
limb of first gill arch; lower jaw not pro- 
jecting. Go to 3. 




3a. Cheek as wide as deep; outline of up- 
per jaw concave. Go to 4. 



3b. Cheek deeper than wide; outline of 
upper jaw not concave. American Shad, 
Alosa sapidissima, page 84, Plate 6. 




Family and Species Accounts 79 



4a. Eye diameter generally less than or 
equal to length of snout; peritoneum 
black. Blueback Herring, Alosa aestivalis, 
below, Plates 4, 5. 



4b. Eye diameter greater than snout 
length; peritoneum pale with dusky 
spots. Alewife, Alosa pseudoharengus, 
page 82, Plate 5. 




Blueback Herring 

Alosa aestivalis (Mitchill 1814) 



Native 
plates 4, 5 




identification. The Blueback Herring can be distinguished from the 
larger shad by the shape of the lower jaw and depth of the cheek (see key 
Figure 3b). It is similar to the Alewife, but the diameter of the Blueback's 
eye is less than or equal to the length of the snout and the peritoneal lining 
of the body cavity is dusky-grey to black. The Blueback Herring's back and 
upper sides tend to be a bluish color. 

selected counts. D 15-20; A 16-21; Scales 41-48; GR (lower) 41-52. 

size. Adults are usually 10 to 12 inches TL. Young-of-the-year are generally 
less than 3 inches TL while in freshwater. 

natural history. Blueback Herring are anadromous; they begin their 
local spawning runs in mid- to late spring when the water temperature 



80 Inland Fishes of Massachusetts 







Blueback Herring: coastal records are not indicated. 

reaches 57°F. Peak spawning occurs with water 
temperature of 69° to 75°F. Spawning occurs in 
swift-flowing sections of streams with gravel or rocky 
bottoms. The fertilized eggs sink to the bottom and hatch 
in less than one week. As in other herring species, fecundity is high with 
each female carrying up to 400,000 eggs. The adults migrate back to salt wa- 
ter after the brief spawning period. The young form large schools and 
slowly work their way downstream to the sea between September and early 
November. Downstream migration is triggered when water temperature 
drops to about 69°F. In freshwater, young Bluebacks eat copepods and 
some cladocerans. In marine waters, adults feed on a variety of marine in- 
vertebrates, including pelagic shrimp. Their first spawning migration oc- 
curs at two to four years. Blueback Herring frequently live to eight years. 



.. /* 



■V.. 



distribution and abundance. Blueback Herring are common in 
Massachusetts and enter numerous coastal streams. Since they were often 
confused with Alewives, little information is available regarding their his- 
torical abundance. However, like other river herrings, their populations 
have been reduced or eliminated in some areas by damming and pollution. 
Bluebacks are abundant in the Connecticut and Merrimack rivers, where 
they migrate as far upstream as New Hampshire or Vermont. More than 
440,000 are passed most years at the Holyoke Fish Lift on the Connecticut 
River (see Figure 3). 

notes. As is true of many other anadromous species, the long migrations 
undertaken by Blueback Herring are currently possible only because of the 
improvements to Massachusetts fishways since the mid-1950s. Besides 



Family and Species Accounts 81 



allowing the fishes simply to pass, the fishways greatly increase the area 
available to Blueback Herring for spawning and feeding (Figure 3). 

references. Bigelow and Schroeder 1953 (general); Clayton etal. 1978 
(review); Domermuth 1976 (food); Scherer 1972 (biology, Connecticut 
River); O'Leary and Kynard 1986 (behavior). 



Alewife 

Alosa pseudoharengus (Wilson 1811) 



Native 
plate 5 




identification. Alewives can be distinguished from the generally larger 
shad by the shape of their lower jaw and the depth of the cheek (key Figure 
3b). Alewives and Blueback Herring are similar but the diameter of an 
Alewife 's eye is greater than the length of the snout, and Alewives have a 
pale peritoneum with small spots that is never dusky to black. The back and 
upper sides tend to be greenish. 

selected counts. D 15-19; A 15-18; Scales 42-50; GR (lower) 36-43. 

size. Adults are usually 10 to 12 inches TL. Young-of-the-year return to the 
sea before they are 4 inches TL. 

natural history. Alewives are anadromous; they spend most of their 
adult life in coastal marine waters and return to freshwaters to spawn. 
Spawning runs begin in midspring as water temperature reaches 52°F. 
During these runs, schools of Alewives swim upstream, spawn numerous 
times over several days, and swim downstream, often passing other schools 
on their way up to the spawning grounds. Spawning occurs in sluggish 



82 Inland Fishes of Massachusetts 



-L'- 




Alewife: open circles indicate stocked populations; 
coastal records are not indicated. 



backwaters of rivers and in ponds. Alewives can 
often be observed spawning in the shallows because 
their splashing makes them obvious. Females release 
60,000 to 300,000 eggs, which stick to the substrate or vegetation. Although 
these annual spawning migrations are physiologically stressful, most adults 
survive and are able to repeat the process in subsequent years. After hatch- 
ing, juveniles form large schools and slowly work their way downstream to 
the sea. While in freshwater, the young feed primarily on zooplankton. After 
reaching marine waters, Alewives feed on zooplankton, small fishes, and 
crustaceans. They become sexually mature after three years and frequently 
live to nine years. 



distribution and abundance. In Massachusetts, Alewives are now 
found in most coastal rivers. Colonial accounts mention their extreme 
abundance. Alewives are still common in some areas, but they have been 
eliminated or reduced in others by damming, pollution, and development. 
Fishways, in place on many streams for hundreds of years, have maintained 
or enhanced numerous populations. Alewives are frequently found in the 
coastal salt ponds on Nantucket and Martha's Vineyard when inlets to these 
ponds have been opened to the sea. They have also been introduced to a 
number of inland lakes in Massachusetts, including Congamond, Single- 
tary, and Webster lakes, and South Pond, Brookfield. 

notes. Alewives are harvested while at sea for a variety of commercial 
purposes, including consumption by humans and other animals. During 
the spawning runs, many fish are dipnetted under town permits in Massa- 



Family and Species Accounts 83 



chusetts. As herrings are often visible and vulnerable during their spawning 
runs, the fishery is strictly regulated to protect this economically important 
species. 

references. Bigelow and Schroeder 1953; Grosslein and Azarovitz 1982 
(general); Clayton et al. 1978 (biology, MA); Belding 1921, Reback and Di- 
Carlo 1972 (distribution, MA); Palmer 1999 (Neponset River). 



American Shad 

Alosa sapidissima (Wilson 1811) 



Native 
plate 6 




identification. American Shad have a deeper than wide cheek and a 
straight upper edge of the lower jaw (see key Figure 3a). As adults, they 
grow to a larger size than any other Massachusetts herrings and have more 
gill rakers than all of the other herrings except the Gizzard Shad. Hickory 
Shad, which are known from Massachusetts marine waters, have a strongly 
projecting lower jaw and only 18-23 lower gill rakers. 

selected counts. D 15-20; A 19-23; Scales 50-55; GR (lower) 59-75. 

size . Most adult American Shad range from 1.5 to 2 feet TL, but they can 
grow larger. The Massachusetts angling record is an 11 -pound, 4-ounce fish 
taken from the Connecticut River in 1986. 

natural history. American Shad are anadromous; they begin their 
migration from the sea to the freshwater spawning grounds in late spring 
when river temperatures reach 50° to 55°F. They may move long distances 
up rivers and migrate into New Hampshire in both the Connecticut and 
Merrimack rivers. Spawning generally occurs in the late afternoon or eve- 



84 Inland Fishes of Massachusetts 



o 




H,u 



y >> 







American Shad: open circles indicate historic records V 4// "U 
where the species may no longer occur; coastal 
records are not indicated. 

ning over shallow areas with sand or gravel sub- 
strates. Females may carry enormous numbers of eggs, 
up to 500,000 in large individuals. After spawning, adult American Shad 
migrate back to marine environments. The eggs gently sink to the bottom 
and roll downstream. Eggs hatch in three to eight days. The young form 
large schools and feed in the river until they grow to about four inches. The 
downstream, seaward- movement of the young is triggered as water temper- 
atures drop to about 66°F. Migration takes place primarily in the late after- 
noon and evening from September to early November. 

Adult American Shad eat a wide variety of zooplankton, shrimp, and 
small fishes. In freshwater, the adults eat little and only occasionally feed 
on small prey. The young-of-the-year feed on small midwater copepods, 
ostracods, and insects. American Shad first spawn at the age of four or five 
years, and adults may live to 10 years of age. 






/.'•' 



-v; 



s 

i /„■■' 



distribution and abundance. Historically in Massachusetts, the 
American Shad entered most coastal streams. Damming, dredging, pollu- 
tion, and other alterations of Massachusetts waters caused large declines 
in the mid- 1800s, when American Shad were eliminated from the Massa- 
chusetts portions of the Connecticut, Blackstone, and Charles rivers. The 
Merrimack suffered declines because fishes were not able to move above 
Lawrence and Lowell. Since the mid-1950s, with new or improved fishways 
and fishlifts, shad numbers have increased dramatically, especially in the 
Connecticut and Merrimack rivers. In many years, nearly 400,000 fish have 
been passed at the Holyoke Fish Lift on the Connecticut River (see Figure 3). 
The species was apparently extirpated from the Blackstone Drainage in the 



Family and Species Accounts 85 



mid- 1800s. Reintroductions made during the late 1970s in the Charles River 
have had minimal success. 

notes. American Shad are a commercially important species, with many 
tons netted in marine waters annually. In freshwater, shad are a popular 
sport fish. The recent recovery of shad in the Connecticut Basin has also 
benefited the Alewife Floater, Anodonta implicata. This freshwater mussel, 
which attaches to the gills of herrings as a larvae, was known only from be- 
low Hartford, Connecticut, before 1970. However, by 1984 and coincidental 
with the shad's range extension, it was found as far north as Bellows Falls, 
Vermont. 

references. Bigelow and Schroeder 1953, Clayton etal. 1978 (review); 
Stevenson 1899, Belding 1921, Reback and DiCarlo 1972, Grosslein and 
Azarovitz 1982 (distribution); Smith 1985 (freshwater mussel); Whitehead 
1985 (review); O'Leary and Kynard 1986 (behavior, migration); Meyer 1999 
(recovery and declines). 



Gizzard Shad Native 

Dorosoma cepedianum (Lesueur 1818) plate 7 




identification. Gizzard Shad have an elongated last dorsal fin ray and 
a slightly inferior mouth, which distinguishes them from all other Massa- 
chusetts herrings. This species also has more than 100 gill rakers, while 
other local herrings have fewer than 75. 

selected counts. D 10-13; A 24-36; Scales 52-70; GR (total) 100-400 
that increase in number with age. 



86 Inland Fishes of Massachusetts 




Gizzard Shad. 



f nP VT 

' & v ( 

size. Gizzard Shad grow 1 8 inches TL, but most \ , / i f ' 
adults are about 12 inches. 



\v 



natural history. Gizzard Shad are primarily a fresh- 
water species, but they may occasionally be found in marine environments. 
This shad lives in a wide range of habitats, including large rivers, swamps, 
reservoirs, canals, and estuaries. In these varied habitats, it generally swims 
in midwater and is usually found in quiet areas with low current. Spawning 
occurs in mid- to late spring, apparently in streams or over shallow bars in 
lakes. Gizzard Shad spawn in groups near the surface, and the fertilized eggs 
slowly sink and stick to the substrate or other underwater objects. As in other 
herring, female Gizzard Shad carry large numbers of eggs; over 400,000 have 
been recorded from large individuals. Gizzard Shad feed almost exclusively 
on phytoplankton, which they filter out of midwater using their numerous 
gill rakers. Juveniles include some zooplankton in their diet. The Gizzard 
Shad is well named because it has a muscular, thick- walled stomach that 
processes food much like the gizzard of a bird. 






distribution and abundance. Gizzard Shad are found over much of 
middle North America west of the Hudson River. In Massachusetts, Gizzard 
Shad were first discovered in May and June of 1985 and 1986, when over 70 
(14 to 18 inches TL) were observed at the Holyoke Fish Lift on the Connecti- 
cut River. Reproduction was confirmed in July 1986 when a 2-inch TL juve- 
nile was collected in the Easthampton Oxbow of the Connecticut River. 
Since 1987, numbers have increased at the Holyoke Fish Lift: 95 (1988); 294 
(1989); 950 (1990); 486 (1991); to 2,065 in 1995. By the early summer of 2000 
the population had exploded and 32,000 gizzard shad were counted as they 



Family and Species Accounts 87 



passed dams on the Connecticut River. In addition, 10 or so specimens 
have been taken each year at the Merrimack River fishways since the first 
was caught in October 1985 (J. O'Leary, pers. comm.). We have seen a pho- 
tograph of an adult taken on hook and line from Mashpee- Wakeby Pond in 
the spring of 1989. In 1991, an adult was found in the Taunton River above 
the Berkley Bridge; in 1997, young were also seen in these waters (S. Hurley, 
pers. comm.). 

notes . The Gizzard Shad is the only freshwater fish species that has natu- 
rally expanded its range into Massachusetts in recent years. Gizzard Shad 
were found for the first time in tributaries to Long Island Sound in the early 
1970s. By 1976, commercial fishermen were catching them near the mouth 
of the Connecticut River, and they were collected 16 miles up the Connecti- 
cut River in 1984. Massachusetts populations are believed to have origi- 
nated from the Hudson River estuary. 

references. O'Leary and Smith 1987 (Massachusetts); Buerkett and Ky- 
nard 1993 (Taunton records); Miller 1960 (systematics and biology); Scott 
and Crossman 1973, Whitehead 1985 (general reviews). 



88 Inland Fishes of Massachusetts 



Anchovy Family 

Engraulidae 



Anchovies are members of the Clupeiformes, or herringlike fishes, and are 
most easily distinguished from the true herrings by the presence of an over- 
hanging snout and a long upper jaw that usually reaches well behind the 
eye. The anchovy family, with about 140 species, has a worldwide distribu- 
tion in temperate and tropical inshore marine environments. Many an- 
chovies can tolerate a wide range of salinities and enter the brackish or 
freshwaters of coastal rivers and streams. A number of tropical species, es- 
pecially in the Amazon Basin, live their entire lives in freshwater. Anchovies 
form an important worldwide fishery, with from 4 to 13 million tons taken 
annually. However, some populations have crashed due to overfishing and 
climatic changes. Decreases in the abundance of anchovies off Peru drasti- 
cally affected the local fish- eating birds and the human economy. An- 
chovies are harvested for human and animal consumption, bait, and a wide 
variety of other uses. 

references. Hildebrand 1963, Whitehead et al. 1988 (reviews). 



Bay Anchovy Native 

Anchoa mitchilli (Valenciennes 1848) 




identification. Anchovies resemble elongated small herrings; however, 
the mouth of the anchovy is large, subterminal, and located well under a 
projecting snout. The upper jaw reaches almost to the posterior edge of the 
gill cover, and the anal fin starts almost directly under the beginning of the 
dorsal fin. Two other species of anchovies, the Striped Anchovy Anchoa 
hepsetus and the Silver Anchovy Engraulis eurystole, have been found in 

Family and Species Accounts 89 




\<i\w v* 



Bay Anchovy. 



1 rV 

)nJJ 



/*> 



= |/V 



Massachusetts marine and brackish waters. The \ ,7 i f 

Bay Anchovy can be easily distinguished from them 

because its anal fin starts almost directly under the be- ^k.<~* 

ginning of the dorsal fin, whereas the anal fin of the other 

two species originates under the end of the dorsal fin (or well behind) 






V :; 



\ c 



% 



*<s, 



selected counts. D 14-16; A 23-30; GR (lower) 20-26. 

size. Adults are usually less than 4 inches TL. 

natural history. The Bay Anchovy is primarily a marine species, but it 
moves seasonally into estuaries and bays to spawn. Where abundant south 
of Cape Cod, they form large schools and are important food for many of 
the larger fishes, such as the Striped Bass and Bluefish Pomatomus saltatrix. 
Anchovies are mostly filter feeders and strain a variety of zooplankton from 
the water with their gill rakers. Spawning occurs from late spring though 
summer; eggs have been found in Cape Cod Bay from June through August. 
The eggs and larvae are pelagic. 

distribution and abundance. The Bay Anchovy is found in Atlantic 
coastal waters from Maine to southern Mexico. In Massachusetts, it is rare 
north of Cape Cod, but small numbers of this anchovy periodically enter 
many of the estuaries south of Cape Cod. During visits to the upper estuar- 
ies, they occasionally enter freshwater, usually during the late summer and 
early fall. Bay Anchovies are never as abundant in Massachusetts waters as 
they are farther south where their biomass may be higher than that of any 
other fish species. 



90 Inland Fishes of Massachusetts 



references. Bigelow and Schroeder 1953, Hildebrand 1963, Whitehead 
et al. 1988 (identification, systematics, natural history); Collette and Hartel 
1988 (Mass. Bay records); Scherer 1984 (eggs and larvae in Cape Cod Bay); 
Vouglitois et al. 1987 (life history, populations). 



Family and Species Accounts 9 1 



Carp and Minnow Family 

Cyprinidae 



The true minnows are found almost worldwide in temperate and tropical 
regions but are absent from South America, Australia, and Madagascar. 
Cyprinids belong to the group of fishes called the Ostariophysi (which in- 
cludes characins, catfishes, suckers, loaches, and electric eels) that have a 
specialized modification of the four or five anterior vertebrae. This modifi- 
cation, called the Weberian Apparatus, links the ear to the swim bladder 
and is used to amplify sound. The Cyprinidae contains more species than 
any other fish family: some 210 genera and more than 2,000 species. As 
adults, they range in size from 1 inch to over 9 feet. The 230 species found 
in North America have many life history strategies, from carnivore to mud- 
eating detritivore, and they live in the slowest rivers and largest lakes to the 
fastest hill- stream torrents. 

Identification of minnows, especially juveniles, may be difficult and great 
care must be taken. Additional confusion results from the many common 
names applied to them. Chub, shiner, dace, and minnow are often used 
interchangeably, and sometimes the terms are used to refer to any of the 
small, bait-sized fishes of other families. The true minnows, however, can 
be identified by the lack of teeth in their oral jaws, their well-developed 
pharyngeal teeth, and one dorsal fin. Male minnows develop contact organs 
or tubercles during the breeding season. These hardened, pointed structures 
are most often found on the snout and pectoral fins. They vary in size and 
placement among species and are used during breeding for territorial ag- 
gression or sexual stimuli. 

Minnows are well known to both anglers and aquarists who use them as 
bait or keep them as pets. Millions of bait-minnows are raised and sold each 
year, and millions of aquarium minnows, such as barbs and rasboras, can 
be found in pet stores. As noted in the following accounts, at least three 
non-native minnow species have been established in Massachusetts 
through bait-bucket introductions, including the Bluntnose Minnow, Fat- 
head Minnow, and Rudd. In addition, the Emerald Shiner, Notropis atheri- 
noides, and the Grass Carp, Ctenopharyngodon idella, have been used as 
live bait or documented from the wild but are not reproducing in the state. 
In the fall of 1997, the Cutlips Minnow, Exoglossum maxillingua, was found 



92 Inland Fishes of Massachusetts 



in the Farmington River and then proven to be reproducing in late 2001. 
Species accounts for these three species are not included in this book, but 
they can be identified by using the keys. 

references. Cavender and Coburn 1992, Fink and Fink 1981, Coburn 
and Cavender 1992, Mayden 1989 (relationships); Winfield and Nelson 1991 
(overview). 



Key to Massachusetts Carp and Minnows 



la. Dorsal and anal fins each with a stout 
serrated "spine" at anterior edge; more 
than 15 dorsal fin rays. Go to 2. 



lb. Dorsal and anal fins without a stout 
serrated "spine" at anterior edge; fewer 
than 11 dorsal fin rays. Go to 3. 




2a. Upper jaw with two fleshy barbels on 
each side; lateral line scales more than 
35 (scales few or absent in "Leather" or 
"Mirror" forms of this species); pharyn- 
geal teeth large and flattened and in 
3 rows. Common Carp, Cyprinus carpio, 
page 104, Platell. 



2b. Upper jaw without fleshy barbels; 
lateral line scales less than 30; pharyn- 
geal teeth not flattened and in 1 row. 
Goldfish, Carassius auratus, page 100, 
Plate 10. 




Family and Species Accounts 93 



3a. Usually over 11 anal rays; deep-bod- 
ied and slab-sided as adults; lateral line 
deeply decurved following ventral out- 
line of body. Go to 4. 




3b. Usually fewer than 10 anal rays (ex- 
cept in Emerald Shiner, Notropis atheri- 
noides, which has 10 to 11); body elon- 
gate and usually not slab-sided; lateral 
line never deeply decurved. Go to 5. 




4a. Belly between pelvic and anal fins 
with fleshy, scaleless keel; 18 to 22 gill 
rakers; 9 to 12 scales above lateral line; 
pharyngeal teeth in 1 row (0,5-5,0). 
Golden Shiner, Notemigonus crysoleucas, 
page 110, Plate 9. 



4b. Belly behind pelvic fins fully scaled, 
without fleshy keel; 10 to 13 gill rakers; 7 
to 8 (seldom 9) scales above lateral line; 
pharyngeal teeth in two rows (3,5-5,3). 
Rudd, Scardinius erythrophthalmus, page 
128, Plate 8. 




5a. Lower lip divided into three lobes, 
the middle tonguelike. Cutlips Minnow, 
Exoglossum maxillingua. See family ac- 
count (not illustrated). 

5b. Lower lip normal, not as above (not 
illustrated). Go to 6. 



94 Inland Fishes of Massachusetts 



6a. Scales small, barely visible, 75 or 
more in lateral line; body with 2 dark lat- 
eral stripes. Northern Redbelly Dace, 
Phoxinus eos, page 118, Plate 12. 

6b. Scales 65 or fewer in lateral line se- 
ries; body with either one dark stripe on 
side or none. Go to 7. 




7a. Anal fin set far back on body, a goes 
into b more than 3 times. Grass Carp, 
Ctenopharyngodon idella. See family 
account. 



7b. Anal origin close to or under dorsal 
fin, a goes into b 2.5 times or fewer. 
Go to 8. 




8a. First dorsal ray short, slightly thick- 
ened and separate from first principal ray 
(in adults); predorsal area tends to be 
flattened, and the scales are small and 
crowded. Go to 9. 




8b. First dorsal ray thin and tightly 
bound to first principal ray; body usually 
rounded and scales moderately crowded 
or well spaced out. Go to 10. 




Family and Species Accounts 95 



9a. Mouth nearly horizontal and over- 
hung by snout; lateral line complete, ex- 
tending to base of tail; a pigment spot on 
anterior dorsal fin rays and at base of 
caudal fin. Bluntnose Minnow, 
Pimephales notatus, page 120, Plate 23. 



9b. Mouth oblique and not overhung by 
snout; lateral line incomplete, not ex- 
tending to base of tail; no noticeable pig- 
ment markings. Fathead Minnow, 
Pimephales promelas, page 122, Plate 24. 





10a. Barbel present either at corner of 
mouth or in groove behind maxilla (not 
illustrated; see key Figures 11a and lib). 
Goto 11. 



10b. Barbels absent (not illustrated). 
Go to 15. 



11a. Barbel at corner of mouth. 
Go to 12. 




lib. Barbel leaflike and in groove 
behind maxilla. Go to 14. 




96 Inland Fishes of Massachusetts 



12a. Upper jaw protractile, with distinct 
groove between premaxilla and snout. 
Lake Chub, Couesius plumbeus, page 102, 
Plate 13. 



12b. Upper jaw not protractile, no 
groove between premaxilla and snout. 
Go to 13. 




12a 



12b 



13a. Snout projecting beyond mouth; 
eye above highest point of upper jaw. 
Longnose Dace, Rhinichthys cataractae, 
page 126, Plate 22. 



«a*> 



13b. Snout not projecting well beyond 
mouth; eye and highest point of upper 
jaw at about the same level. Blacknose 
Dace, Rhinichthys atratulus, page 124, 
Plate 21. 



/$*(§) 




14a. Lateral line scales fewer than 50; 
most body scales with dense pigment at 
anterior edge; no black spot at leading 
edge of dorsal fin. Fallfish, Semotilus cor- 
porate, page 132, Plate 18. 



14b. Lateral line scales more than 52; 
anterior edge of scales without dark pig- 
ment; dark spot usually present at ante- 
rior base of dorsal fin. Creek Chub, Se- 
motilus atromaculatus, page 130, Plate 19. 




Family and Species Accounts 97 



15a. Lining of body cavity black; intes- 
tine long and coiled; small bump at tip 
of lower jaw. Eastern Silvery Minnow, 
Hybognathus regius, page 106, Plate 14. 



15b. Lining of body cavity silvery, with or 
without dark speckles; intestine short, 
less than twice standard length; no bump 
at tip of lower jaw. Go to 16. 




16a. Anal fin with 10 to 11 rays. Emerald 
Shiner, Notropis atherinoides. See family 
account. 

16b. Anal fin with 7 to 9 rays (not illus- 
trated). Go to 17. 




17a. Scales along anterior portion of lat- 
eral line deeper than wide. Go to 18. 



17b. Scales along anterior portion of lat- 
eral line equally deep as wide. Go to 19. 




98 Inland Fishes of Massachusetts 



18a. Predorsal scales over 22; anal fin 
usually with 9 rays; more than 5 scale 
rows above lateral line; no V-shaped dark 
pigment behind anus; eye diameter less 
than snout length. Common Shiner, 
Luxilus cornutus, page 108, Plate 17. 



18b. Predorsal scales approximately 15; 
anal fin usually with 7 rays; less than 
5 scale rows above lateral line; V-shaped 
area of dark pigment behind anus; eye 
diameter greater than snout length. 
Mimic Shiner, Notropis volucellus, 
page 116, Plate 16. 




19a. Usually 8 anal fin rays; dorsal, anal, 
and pectoral fins typically falcate; no dark 
band along snout and body (except young 
may have a weak band). Spottail Shiner, 
Notropis hudsonius, page 114, Plate 15. 

19b. Seven anal fin rays; dorsal, anal, 
and pectoral fins not falcate; dark band 
through eyes and across snout. Bridle 
Shiner, Notropis bifrenatus, page 112, 
Plate 20. 




Family and Species Accounts 99 



Goldfish 

Carassius auratus (Linnaeus 1758) 



Introduced 
PLATE 10 





3 «*|r; 



identification. Like carp, Goldfish are heavy-bodied minnows with 
large scales, a long dorsal fin, and hardened and serrated anterior anal and 
dorsal rays. Goldfish lack the mouth barbels found in carp, and their pha- 
ryngeal teeth are in one row (see key Figure 2b). The natural goldfish color 
is olive to brassy; however, propagated fish may range from all gold to 
orange or to mixtures of red, white, black, and orange. Introduced popu- 
lations may gradually revert to the natural wild color. 

selected counts. D i,15-19; A i,5-6; Scales 25-31; PT 0,4-4,0. 

size. Goldfish can grow to be a foot or more in length but most Massachu- 
setts specimens are only 5 to 8 inches TL. The largest specimen that we 
have seen is 13 inches TL (215 mm SL) and was collected from the Charles 
River, Cambridge. 

natural history. Introduced Goldfish seem to do best in smaller ponds 
with abundant aquatic vegetation. We have rarely found them in flowing 
waters. Goldfish spawn in the spring (May- June in New York). Two or more 
males follow single females over aquatic vegetation to spawn. The spawn- 
ing behavior is fast and accompanied by aggressive splashing. In fact, it can 
be quite violent; delicate, ornamental strains are often damaged while re- 
producing in aquaria. In the wild, eggs are scattered over the bottom vegeta- 
tion and hatch in three to four days. In most populations, female goldfishes 
are more abundant than males; there are 13 to 36 males reported for each 
100 females. Males are also smaller and grow more slowly than females. 



100 Inland Fishes of Massachusetts 







Goldfish. 



Goldfish mature anywhere from nine months to 

four years depending on the strain and the envi- 

ronment. Adults have been known to live for six or W* 

seven years. Goldfish are omnivorous, consuming a wide 

range of food types including larval and adult aquatic insects, mollusks, 

crustaceans, worms, and vegetation. 









distribution and abundance. Goldfish are native to eastern Siberia, 
China, and Korea, but have been introduced worldwide. Introduced popu- 
lations are now found in every state in the United States, except Alaska, 
and in three Canadian provinces. Goldfish were the first exotic fish to be 
brought to North America. In his 1842 review of New York fishes, J. DeKay 
reports that the first releases were as early as the late 1600s. Goldfish were 
common and well known in the waters around Brookline, Cambridge, and 
Brighton, Massachusetts, before 1839, but the species was not noted in 
western Massachusetts prior to 1941. We have found specimens in scattered 
areas throughout the state, usually near urban centers. The species is prob- 
ably more widely distributed than our data suggest since Goldfish are com- 
mon in farm and golf course ponds, which we did not survey. 

notes. Goldfish and Common Carp hybridize and produce fertile off- 
spring; however, we have not found hybrids in Massachusetts. 

references. Breder and Rosen 1966 (reproduction); Courtenayand 
Stauffer 1984, Storer 1839, DeKay 1842 (introductions); Scott and Crossman 
1973 (Canada); Smith 1985 (NY). 



Family and Species Accounts 101 



Lake Chub Native, State Endangered 

Couesius plumbeus (Agassiz 1 850) plate 13 



identification. Lake Chub are elongate, moderately round-bodied 
minnows with a small but well- developed conical barbel at the posterior 
end of their upper jaw. The snout is completely separated from the upper 
lip by a continuous deep groove, and the mouth is slightly subterminal. 
Breeding males have a hint of orange wash on the pectoral fins and face. 

selected counts. D 8; A 8; Scales 10/53-70/7; PT 2,4-4,2. 

size . Adults are between 3 and 4 inches TL, but specimens up to 9 inches 
TL have been found outside of Massachusetts. 

natural history. Due to its rarity, almost nothing is known about the 
behavior of Lake Chub in Massachusetts. Relatively few studies of Lake 
Chub have been conducted, and those that are available generally pertain 
to lake habitats, which the species prefers. In Massachusetts, the Lake Chub 
has been found only in moderate to fast-flowing, clear, cold streams. It pre- 
fers areas of little or no vegetation with gravel and rubble bottoms. Lake 
Chub spawn during late spring to early summer. In the Connecticut Lakes 
Region of New Hampshire, spawning occurs in early July. Nests are not 
built; eggs are simply deposited on rocky substrate and left unguarded. In 
British Columbia, Lake Chub mature in their third year and seldom live 
more than five years. Females grow faster and live longer than males. Lake 
Chub feed on a variety of stream invertebrates, including aquatic insects 
and crustaceans. Occasionally, they will eat small fishes and algae. 

distribution and abundance. Lake Chub are found throughout 
Canada and at scattered localities in the United States from northern New 



1 02 Inland Fishes of Massachusetts 



<U U\ 







Lake Chub: open circles indicate known localities 
where species was not found during our post- 1969 
surveys. 






\lt\ S< 



i >-■■■■ •-. 

) 9 .. P 



ii\ 



/,■■ 



KjV.*.!*— 



England to New York and Michigan. In Massachu- & 

setts, Lake Chub are rare and are currently uncommon in 
the upper portions of the Westfield River. As late as 1952, Lake Chub were 
common in the Middle and West branches of the Westfield; however, sur- 
veys conducted between 1977 and 1990 have failed to locate this species in 
the Middle Branch and have found only a few specimens in the upper East 
and West branches. The Westfield population is disjunct; the nearest popu- 
lation is in the northern Connecticut River Basin of Vermont and New 
Hampshire. 



notes. The Massachusetts population occupies the southeastern-most 
part of the species' range. Lake Chub populations at the southern extremes 
of its range are often disjunct, and several of these populations may have 
been extirpated. Lake Chub are currently listed as State Endangered by the 
Massachusetts Division of Fisheries and Wildlife. Their listing is due to a 
documented decline over the last 30 years. This species has been collected 
at only a few of the many sites surveyed in the Westfield Drainage since 
1977. The reasons for its decline are unknown. 

references. Brown et al. 1970 (breeding); Halliwell 1978 and 1989 (sur- 
veys); McCabe 1942, 1943, Mullan 1952 (Westfield records). 



Family and Species Accounts 1 03 



Common Carp Introduced 

Cyprinus carpio Linnaeus 1758 plate i i 




identification. Common Carp are large, robust-bodied minnows with 
long dorsal fins. The first rays of the dorsal and anal fins are modified into 
stout, serrated spines. Two pairs of barbels on the upper jaw separate Com- 
mon Carp from Goldfish. The scale number is variable: in a variety called 
"leather carp," scales maybe absent; in another called "mirror carp," scales 
are enlarged and scattered, but the typical form has about 35 large scales. 
The pharyngeal teeth in carp are heavy and molarlike and more developed 
than in Goldfish. 

selected counts. D i,18-23; A i,4-5; Scales 32-41(0); PT 1,1,3-3,1,1. 

size . Most adult Common Carp are around 2 feet TL, but they can grow 
much larger. A 44.1 pound carp angled from the Connecticut River in 1993 
is the current Massachusetts state record. 

natural history. Common Carp usually inhabit large, slow-flowing 
rivers, large ponds, and lakes with abundant aquatic vegetation. Carp win- 
ter in deep water but move inshore during spring. Spawning carp are often 
seen swimming and rolling with their backs and dorsal fins out of the water 
when water temperatures exceed 59°F, and spawning continues from late 
spring into late summer. Carp spawn in inshore areas with aquatic or sea- 
sonally flooded vegetation. Groups of three to four males spawn with each 
female and their behavior causes splashing and uprooting of vegetation. 
The eggs are randomly broadcast and adhere to vegetation until they hatch 
in 4 to 12 days, depending on temperature. Female carp carry an enormous 
number of eggs, with fecundity increasing with size. Large females (about 



104 Inland Fishes of Massachusetts 




Common Carp. 



w\y 



k 



35 inches TL) carry over 2 million eggs. Juvenile 

carp grow rapidly, reaching 5 to 6 inches in the first 

year of growth. Carp may live at least 20 years and may \ * * 

grow to 60 pounds; however, most adults in a population 

weigh 4 to 15 pounds and are four to eight years old. Carp are omnivorous, 

eating great quantities of animal and vegetable matter. Their diet has been 

found to include leaves, roots, stems and seeds of aquatic plants, seeds of 

terrestrial plants, worms, leeches, crustaceans, mollusks, and occasionally 

fish eggs. 



distribution and abundance. Common Carp are native to almost 
all of Eurasia, but the exact native range is unknown due to pre-Roman in- 
troductions. They were brought to North America as early as 1831 and 
through subsequent introductions and natural dispersal they are now found 
in all of the lower 48 states, Hawaii, and southern Canada. Common Carp 
were first distributed in Massachusetts by the U.S. Bureau of Fisheries in 
1880. Today, Common Carp are found in many areas, particularly the Merri- 
mack, Concord, Connecticut, Taunton, and Blackstone rivers and in a num- 
ber of larger lakes and ponds. Carp are at times common; over 20,000 were 
killed by dropsy (caused by an Aeromonas bacteria) over a short period in 
the Merrimack River in the late 1970s. Our records probably underestimate 
the range and abundance of this species since it is normally not taken with 
the small seines and electrofishing gear used during our surveys. Koi, which 
are often found in garden ponds, are an ornamental variety of Common 
Carp and are not Goldfish. 

notes. It became evident that Common Carp sometimes interacted in a 
negative way with its new environment as early as the turn of the century. 



Family and Species Accounts 105 



Most ecological problems associated with carp center around its alteration 
of habitat during foraging and spawning. Taylor et al. (1984:336) state, "De- 
terioration in native fish populations has often accompanied the spread 
and buildup of carp populations... Evidence for impact is strong in the sense 
that, in numerous independent studies, increases in carp population and 
concomitant changes in habitat structure have been repeatedly associated 
with declines in or displacement of native assemblages. However, the multi- 
plicity of effects possible — given the complex manner in which carp interact 
with virtually every physical and biological component of an ecosystem — 
has made it difficult to pinpoint simple cause-effect relationships." The 
habitat disturbance caused by carp has also contributed to a decline in the 
quality of waterfowl habitat in some areas. However, some authors state 
that the carp is not the "villain" that it has been long labeled (Jenkins and 
Burkhead 1993:275). 

references. Garman 1889, 1890, Courtenay and Stauffer et al. 1984 (in- 
troductions); MacCrimmon 1968, Cooper 1987 (biology); Taylor et al. 1984 
(impacts); Mirick 1991 (MA). 



Eastern Silvery Minnow Native, State Special Concern 

Hybognath us regius Girard 1856 plate 14 




identification. The Eastern Silvery Minnow is a rather stout, round- 
bodied shiner with medium-sized eyes. It is distinguished from other Mas- 
sachusetts minnows by a combination of characteristics: a small, slightly 
subterminal mouth; a lower jaw with a fleshy knob at the tip; a black lin- 
ing of the body cavity (peritoneum); a long, coiled intestine often seen 
through the belly wall; an expanded and flattened posterior extension of 
the skull (the basioccipital process); and 38 to 40 lateral line scales. It is 
silvery all over. 

1 06 Inland Fishes of Massachusetts 




C\-^r 



h-tk^^M^M (4^/" t s \ ,\ \^;y :^t 



Eastern Silvery Minnow. 



U/V NVJ >.s 



selected counts. D 8; A 8-9; Scales 6/38- 
40/4; PT 0,4-4,0. 



size . Adults usually range from 3 to 5 inches TL 




natural history. Eastern Silvery Minnows characteristically inhabit 
wide, slow-moving rivers. They spawn diurnally in late spring at tempera- 
tures of 55° to 69°F in backwaters and lower reaches of tributary streams. 
This minnow is unique among northeastern cyprinids in that it lays nonad- 
hesive eggs directly on bottom ooze in areas where emergent grasses and 
reeds provide cover. Females do not spawn until they are in their second 
year of life. Detailed studies of the diet of this species have not been carried 
out; however, filamentous algae and organic matter filtered from bottom 
ooze constitute the main food sources. Filtering is accomplished by modi- 
fied papillae in the throat, and this filtered material is efficiently processed 
by the long, coiled intestine. 

distribution and abundance. In Massachusetts, this species is 
known only from the main stem of the Connecticut River north of the 
Holyoke Dam and in the lower Deerfield River. During the 1950s, Prof. 
Thomas J. Andrews found that this species was common over the flooded 
flats along the Connecticut River near Hadley. One of his seine collections 
contained nearly 100 specimens. However, surveys between 1978 and 1990 
have recorded only a few individuals, usually collected along with the 
abundant Spottail Shiner. 

notes. Since this minnow has apparently declined over the past 30 years, 
it is currently listed as a State Species of Special Concern. The reasons for 



Family and Species Accounts 107 



this decline in Massachusetts are uncertain; however, other members of the 
genus Hybognathus have been noted as declining in the Midwest due to sil- 
tation, pollution, and changes in water flow. In Massachusetts, the decline 
may be related to human manipulation of the natural river flow in the Con- 
necticut Valley, as dams and pump storage facilities have been built. These 
types of water control practices may reduce or change the character of 
backwaters and spawning sites used by this minnow. Until recently, this 
species was considered a subspecies of the Mississippi Silvery Minnow, 
Hybognathus nuchalis. 

references. Hlohowskyj et al. 1989 (filtering apparatus); Pflieger 1975 
(declines, Missouri); Smith 1979, Warren and Burr 1989 (declines, Illinois); 
Raney 1939 (biology). 



Common Shiner 

Luxilus cornutus (Mitchill 1817) 



Native 
plate 17 




identification. Common Shiners are relatively deep-bodied minnows 
with a combination of 9 anal rays (rarely 8 or 10); deeper- than- wide anterior 
lateral scales (see key Figure 17a); and more than five scales above the lat- 
eral line. Common Shiners have distinctive horizontal stripes that appear in 
three bands; a pale middorsal band, a darker stripe below it, and a second 
pale stripe below that. In breeding males the stripes become golden and the 
body bronze; dark crescent-shaped marks appear on the body; the head 
darkens to blue -gray; and the fins darken with a pink to red distal edge. 

selected counts. D 8; A 9 (8-10); Scales 7-8/38-44/5; PT 2,4-4,2. 

size . Common Shiners are a medium to large minnow, often reaching 5 to 
6 inches TL; some Massachusetts specimens reach 7 inches TL (135 mm SL). 



108 Inland Fishes of Massachusetts 



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Common Shiner: open circles indicate known locali- f { // "i . - ^% s SKlJ\ > 
ties where species was not found during our post- 1969 V// ^ y )M „ 

surveys; not all solid circles were resurveyed. 7 V / 



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natural history. In Massachusetts, Common v a 

■ 

Shiners are most often found in large rivers to small 
streams with relatively clean water. Upstream spawning migrations begin 
in May as water temperatures reach 60° to 65°F. Males, which are much 
larger than females, establish and defend territories. Spawning sites are 
usually over gravel beds in running water where males sometimes excavate 
small depressions or use the spawning sites of other nest-building min- 
nows. In Massachusetts, these nest builders include Fallfish and Creek 
Chub. This communal spawning behavior may result in hybrids between 
the Common Shiner and other species. Occasionally, up to 100 males may 
gather at a nest, and there is constant jostling for optimal positions on the 
site. The spawning act takes a fraction of a second, after which the partici- 
pants drop downstream. Males and females may return many times to 
spawn with the same or different partners. Probably fewer than 50 eggs are 
laid at each spawning. Common Shiners feed mainly at the surface or in 
midwater, but they are opportunistic feeders. Aquatic insects, including 
both adults and larvae, are the primary food source, but small fishes and 
some plant material are also eaten occasionally. 

distribution and abundance. In Massachusetts, this minnow is 
most common from the Connecticut Drainage west, where it is found in all 
of the major Connecticut River tributaries and in the Hoosic and Housa- 
tonic rivers. In addition, there are scattered records from the Nashua, Mer- 
rimack, French, Blackstone, Taunton, and Charles river drainages. It is ab- 
sent from all coastal streams, Cape Cod, and the Islands. As noted, the 
species may be declining. 






Family and Species Accounts 109 



notes. Common Shiners may have been more widely distributed in east- 
ern Massachusetts in the past. For instance, we have seen historic speci- 
mens from the Charles Drainage collected at Waltham (late 1800s) and at 
Medfield (1962), but we have not observed this species in the Charles Drain- 
age during any of our post- 1975 surveys. Other eastern Massachusetts rec- 
ords, except from the Merrimack River Drainage, are scattered and rare. A 
recent comparison of the results of pre- 1950 stream surveys to post- 1975 
surveys shows a considerable decline in the relative occurrence of the 
Common Shiner in the central portions of the state, particularly in the 
Millers and Chicopee drainages. Similar declines in this species have been 
noted within its midwestern range. This species was formerly placed in the 
genus Notropis. 

references. Gilbert 1964 (description, distribution, relationships); Halli- 
well 1989 (declines, MA); Raney 1940a (breeding). 



Golden Shiner Native 

Notemigonus crysoleucas (Mitchill 1814) . plate 9 




identification. Golden Shiners are deep-bodied, compressed fish with 
a down-curved lateral line. Among North American minnows, it is unique 
in having a fleshy, scaleless area on the ventral midline between the pelvic 
fins and the anus. This characteristic is often difficult to see in small speci- 
mens. The species can also be confirmed by the long anal fin, which usually 
has over 12 rays. Adults are brassy with orange fins. The Golden Shiner is 
similar to the introduced Rudd, which lacks the fleshy keel and has larger 
scales, fewer gill rakers, and two rows of pharyngeal teeth. Juveniles might 
be confused with other minnows because they are not deep-bodied and 



1 10 Inland Fishes of Massachusetts 




Golden Shiner: open circles indicate known localities 
where species was not found during our post- 1969 
surveys; not all solid circles were resurveyed. 

have a dark band on the body, but the anal count 
will separate the species in most cases. 



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selected counts. D 7-9; A 10-15; Scales 9-12/41-50/3-4; GR 18-22; 
PT 0,5-5,0. 

size . This minnow commonly reaches 8 or 9 inches TL, but specimens 
close to 12 inches TL have been reported. 

natural history. Golden Shiners are found in a wide range of habitats, 
including lakes, ponds, and slow-moving rivers and streams. They spawn in 
spring and summer, from May to August. Spawning begins when the water 
temperature is around 70°F. The adhesive eggs are broadcast over sub- 
merged vegetation in shallow water, and the adults do not guard or other- 
wise tend the eggs. The young grow fast during their first summer and may 
reach 2 to 3 inches by fall. Most Golden Shiners do not spawn until their 
third summer and carry up to 200,000 eggs. Golden Shiners are midwater 
and surface feeders, often picking individual small prey out of the water 
column. They feed mainly on zooplankton, but adults sometimes feed on 
insects and small fishes. Algae are also an important part of their diet. 

distribution and abundance. In Massachusetts, Golden Shiners are 
abundant and widely distributed. This species occurs in every drainage in 
the state, and its distribution has been enhanced by the release of fishes 
from bait-buckets. While the largest specimens are found in slow back- 



Family and Species Accounts 111 



waters of rivers and large ponds, small juveniles are often found in smaller 
hill streams. 

notes. Golden Shiners, sometimes called "pond shiners," are an impor- 
tant forage species for game fishes. They are the most common bait fish 
sold in Massachusetts, and many are imported into the state from areas 
where they are commercially propagated. The Golden Shiner is known to 
hybridize with the introduced Rudd. 

references. Keast and Webb 1966 (feeding ecology); Scott and Cross- 
man 1973 (biology, variations, Canada); Burkhead and Williams 1991 (hy- 
brids, identification); Smith 1985, Jenkins and Burkhead 1993 (general). 



Bridle Shiner Native, State Special Concern 

Notropis bifrenatus (Cope 1869) plate 20 




identification. Bridle Shiners are small minnows with a distinct dark 
lateral band that runs forward through the eye and around the snout; a 
small mouth; seven anal rays; and an incomplete lateral line. The young of 
other minnow species and young chubsuckers sometimes show a dark lat- 
eral band(s) but do not have the Bridle Shiner's large, outlined scales on the 
lateral body. These scale outlines are noticeable in specimens less than 
1 inch TL (22 mm SL). Adults are straw-colored, and breeding males de- 
velop an intense yellow wash along the sides. 

selected counts. D 8; A 7; Scales 4-5/32-36/4; PT 0,4-4,0. 

size . This is a small minnow, usually under 2 inches TL. The largest speci- 
men we have seen measured just over 2.25 inches TL (46 mm SL). 



1 12 Inland Fishes of Massachusetts 




Bridle Shiner: open symbols indicate localities 
where species was not found during our post- 1969 
surveys (circles) or during the 1993-1995 New England 
Aquarium resurvey (squares) of known localities. 






i 






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natural history. The Bridle Shiner is typically 
found in well-vegetated, quiet waters where schools often swim in and out 
of vegetation along the edge of ponds. Breeding occurs from late May to 
mid-July, and sometimes into August. Males usually pursue the larger fe- 
males just below the surface in areas of open water over submerged vegeta- 
tion. Spawning always occurs near the surface. About 10 eggs are released 
at each mating, and they fall to the bottom. Spawning is repeated many 
times. In an aquarium, over 320 eggs were released in a two-hour period. 
Eggs hatch in two to three days at 75°F, and by six weeks, the young resemble 
the adults. Bridle Shiners feed almost exclusively on animal matter, includ- 
ing small aquatic insects, copepods, cladocerans, and ostracods. This spe- 
cies is short-lived; most adults die during their second year. Bridle Shiners 
are prey for many larger fishes, particularly pickerel and bass. 



■-?.'■/ 



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distribution and abundance. In Massachusetts, we have examined 
Bridle Shiner specimens from all major river basins except the Islands. 
Jenkins and Zorach, in their 1970 distributional study, noted that the Bridle 
Shiner was excluded from the upper Connecticut River by the fall line at 
Turners Falls. We, however, have collected one specimen from Willow 
Brook, a tributary to South Athol Pond. This single Millers Drainage record 
is probably an introduction or due to stream capture with the nearby 
Quabbin watershed. 

note . Although Bridle Shiners were common at least until the early 1960s, 
this interesting little minnow is currently declining in eastern Massachu- 



Family and Species Accounts 113 



setts, where few specimens were collected in our surveys between 1975 and 
1989. A New England Aquarium survey (1993-1995) of many known locali- 
ties found the Bridle Shiner at only 23 percent of its former sites in eastern 
Massachusetts. This shiner has a relatively small range, from southern New 
England to South Carolina, and it has been extirpated or is declining in 
much of the region. The Bridle Shiner is listed as a Species of Special Con- 
cern in Massachusetts. Recently it has been suggested that the Bridle 
Shiner belongs in the genus Hybopsis and not Notropis. 

references. Burkhead and Jenkins 1991, Whittier et al. 1997, Sabo 2000 
(declines); Harrington 1947a (development), 1947b (breeding), 1948a (life 
cycle), 1948b (food), 1951 (spawning); Jenkins andZorach 1970 (zoogeogra- 
phy and morphology), Mayden 1989 (placement in Hybopsis). 



Spottail Shiner Native 

Notropis hudsonius Clinton 1824 plate 15 




identification. Spottail Shiners are medium- sized silvery minnows 
that usually lack obvious pigment patterns except for a diffuse midlateral 
stripe and a spot at the base of the caudal peduncle. This spot is often ob- 
scured in larger specimens. The mouth is slightly subterminal, median fins 
are often falcate, and the anterior lateral line scales are as wide as they are 
deep (see key Figure 17b). 

selected counts. D 8; A 7-8; Scales 5/36-43/4-5; PT variable 0,4-4,0 
to 2,4-4,2. 

size . Adults are normally about 4 inches TL; however, a few specimens 
over 5 inches TL (1 10 mm SL) have been found in Massachusetts. 



1 14 Inland Fishes of Massachusetts 




K 



Spottail Shiner. 

natural history. Spottail Shiners are found 

primarily in larger rivers and only occasionally in 

large reservoirs and lakes. Spottail Shiners spawn from . ;l|W^ 

May to mid-June, with the onset of reproductive activities 

apparently tied to water temperature. Although their spawning behavior 

has not been extensively studied, Spottail Shiners form large aggregations 

and scatter their eggs on sandy bottoms at the mouths of streams. Large 

females contain up to 2,700 eggs but, as in many other fishes, fecundity is 

related to size. The young grow fast and often reach 2 to 3 inches TL by the 

end of their first year. Growth slows after the first year, and it usually takes 

another two years for individuals to reach 4 inches TL. The species may live 

five years. Spottail Shiners tend to feed near the bottom and consume small 

mollusks, mayflies, and other aquatic or terrestrial insects. Adults also feed 

on large numbers offish eggs, including their own. 



distribution and abundance. In Massachusetts, this minnow is 
abundant in the Connecticut, Deerfield, Chicopee, and Westfield drainages. 
It is common in the Merrimack and Housatonic river drainages, and a few 
specimens have been collected from the Neponset, Nashua, and Concord 
rivers. During the late 1970s, Spottail Shiners were common in the lower 
Charles River in Cambridge and Boston, but we have taken only a few spec- 
imens since 1985. Steven Shapiro (1976), who studied the species, thought 
that the Massachusetts populations outside of the Connecticut Basin most 
likely resulted from bait fish introductions. This may be true since they are 
absent from the Blackstone and Taunton river drainages, where this species 
might be expected to occur, and Spottail Shiners were not mentioned by 
early authors such as Storer, Putnam, and Goode and Bean. 



Family and Species Accounts 115 



notes. Over most of their range, Spottail Shiners are considered an im- 
portant forage and bait minnow. 

references. Shapiro 1975 (bibliography), 1976 (age and growth, diet); 
Wells and House 1974 (life history). 



Mimic Shiner introduced 

Notropis volucellus (Cope 1865) plate i 6 



identification. Mimic Shiners are a large-eyed, silvery minnow best 
identified by examining scales and pigment. Anterior lateral-line scales are 
deeper than they are wide (see key Figure 17a), lightly outlined, and large 
(fewer than 15 predorsal scales and fewer than five scale rows above the lat- 
eral line). A V-shaped pigment spot behind the anus, in combination with 
some pigment along the base of the anal fin, is also characteristic. In addi- 
tion, a spot at the caudal base is expanded into a diffuse oval with a small 
forward-facing triangle at the base of the middle caudal rays. 

selected counts. D 8; A 8; Scales 4/33-38/3-4; PT 0,4-4,0. 

size . Mimic Shiners seldom grow larger than 3 inches; however, we have 
collected a 4-inch TL (96 mm SL) specimen from Massachusetts. 

natural history. Within their native range, Mimic Shiners inhabit 
clear streams, smaller rivers, and lakes. In Massachusetts, they are primar- 
ily found in the main channel and quiet backwaters of large rivers. Almost 
nothing is known about this introduced minnow in Massachusetts, and 
only a few studies have been conducted in its native range. This species ap- 
parently spawns in the summer (June-July) and possibly at night. Eggs are 
deposited over aquatic vegetation and are not cared for by the adults. The 



116 Inland Fishes of Massachusetts 



■% V 



PC 




Mimic Shiner. 






Mimic Shiner's food consists mainly of small 
aquatic invertebrates and some algae. This small 
minnow usually does not live more than two years. 




* 






■-'y. 



distribution and abundance. In Massachusetts, Mimic Shiners 
were first found in a small tributary of the Connecticut River near Long- 
meadow by B. McCabe in 1941. Their introduction into Massachusetts 
waters probably resulted from bait fish releases before that date. Today, 
they are common in some areas of the Connecticut main stem and in the 
lower Westfield and Deerfield rivers. A small series of juveniles, collected 
by Prof. T.J. Andrews in 1953 from Townsend Harbor on the Squannacook 
River, is the only known record outside the Connecticut Basin. 

notes . This shiner is native to a wide area west of the Appalachian Moun- 
tains and the Saint Lawrence Drainage. A small Atlantic slope population in 
North Carolina and Virginia is thought to be native. 

references. Black 1945 (life history); Olmsted et al. 1979 (feeding); Scott 
and Crossman 1973 (Canada); Smith 1985, Jenkins and Burkhead 1993 
(general). 



Family and Species Accounts 117 



Northern Redbelly Dace Native, State Endangered 

Phoxinuseos (Cope 1862) plate 12 




identification. Northern Redbelly Dace have two horizontal dark or 
dusky stripes along their upper sides, small scales, a long, coiled intestine, 
and a black lining of the body cavity. The upper band is often broken into 
small dots or patches behind the dorsal fin, but the lower midlateral band 
is always complete. The flanks, belly, and throat range from creamy white 
in immatures to yellow or red in breeding adults. 

selected counts. D 7-8; A 7-8; Scales 70-90; PT 0,5-5,0. 

size. This is a small minnow; most adults are only about 2 inches TL. 

natural history. Little is known about the Massachusetts population 
of this species. However, studies done elsewhere indicate that Northern 
Redbelly Dace breed in early spring to midsummer and that they may 
spawn two times each season. When spawning, a female, accompanied 
by one or more males, darts into a clump of filamentous algae, where 5 to 
30 nonadhesive eggs are released and fertilized. Male Northern Redbelly 
Dace have comblike, tuberculate scales in front of their pectoral fins that 
may be related to reproductive activity. Eggs hatch in 8 to 10 days at 70° to 
80°F, and the young may not reach maturity until their second or third sum- 
mer. When found with Finescale Dace, Phoxinus neogaeus, the Northern 
Redbelly Dace reproduces in an interesting mosaic of diploid and triploid 
hybrids. Northern Redbelly Dace are long-lived for such small fishes; a 
Canadian study showed that some individuals live up to eight years. Like 
many herbivorous animals, Northern Redbelly Dace have long, coiled in- 
testines. These dace feed mainly on plant material, primarily diatoms and 
filamentous algae, but also eat zooplankton and insects. 



118 Inland Fishes of Massachusetts 




Northern Redbelly Dace: open circles indicate 
known localities where species was not found during 
our post- 1969 surveys. 

DISTRIBUTION AND ABUNDANCE . Although 

common in some areas of northern New England, North- 
ern Redbelly Dace are rare in Massachusetts. They are known only from a 
small portion of the Green River system (Deerfield Drainage) in the vicinity 
of Greenfield. The Massachusetts population is historically known from only 
four localities in the Green River. B. McCabe first discovered this species in 
Massachusetts near downtown Greenfield in 1940, but it is no longer found 
at this site. In fact, these dace have been found only in a single small tribu- 
tary since 1978, where they are uncommon. Changing land-use patterns 
and development, resulting in changes in water quality, could easily extir- 
pate this disjunct population. 






y, 



notes. Over most of its range, Northern Redbelly Dace are found in boggy, 
acidic environments, but in Massachusetts they inhabit a nonboggy clear 
stream and associated spring-fed seepage pools. The Massachusetts popu- 
lation is geographically isolated from the nearest New England population 
in the Sugar River system in the Connecticut Basin in New Hampshire. This 
species had been treated as a member of the genus Chrosomus, but recently 
it has been documented that it should be placed in the genus Phoxinus. 
This change reflects a close relationship between the six North American 
species and their Eurasian relatives. 

references. Cochran et al. 1988 (diet); Bailey and Oliver 1939 (NH popu- 
lation); Howes 1985 (systematics and anatomy); McCabe 1942 (first MA 
records); Scott and Crossman 1973 (review, Canada); Goddard et al. 1989, 
Goddard and Schultz 1993 (hybrids). 



Family and Species Accounts 119 



Bluntnose Minnow introduced 

Pimephales notatus (Rafinesque 1820) plate 23 




identification. Bluntnose Minnows are most similar to Fathead Min- 
nows, which also have a well- developed but short first dorsal ray that is 
separated from the first principal ray (in adults) (see key Figure 8a) and 
small, irregular, crowded scales from the nape to the dorsal fin. Its body is 
quite round, almost square in cross section, and is elongate with a com- 
plete lateral line, a cross-hatched scale pattern, and a well-marked spot at 
the base of the caudal fin. This species has a blunt head with a slightly sub- 
terminal mouth. It has a coiled intestine, a dark peritoneum, and a dark 
band along the snout and body. Males become dark, almost black, during 
the breeding season. 

selected counts. D 8; A 7; Scales 6/42-50/4; PT 0,4-4,0. 

size. Adult Bluntnose Minnows typically range from 3 to 4 inches TL. 

natural history. Bluntnose Minnows are found in a wide variety of 
habitats but usually prefer sandy to gravelly substrates. Spawning lasts 
from spring to midsummer. Males excavate a pocket under flat stones, 
wood, or cans to make a nest. Males vigorously guard the nest during the 
spawning season and chase away rival males and all other species of fishes. 
After a female enters a nest, she deposits a number of adhesive eggs on the 
undersurface of the roof. Eggs are added to the nest by a number of females; 
up to 2,500 eggs have been found in a single nest. Spawning is prolonged 
and different- aged eggs may be found in the same nest. Males also clean 
the eggs, remove dead or diseased eggs, and circulate water in the nest. If 
the male is removed from the nest, the eggs will not hatch. Eggs hatch in 
10 to 14 days, and the young reach almost 2 inches TL at the end of their 



120 Inland Fishes of Massachusetts 










> 



Bluntnose Minnow. 



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first growing season. Males are larger than fe- 
males and do not mature until their second year. 
Bluntnose Minnows are primarily bottom feeders and 
often sift through large amounts of bottom material in 
search of a variety of invertebrates. 



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distribution and abundance. Bluntnose Minnows are not native to 
Massachusetts, and the species was not found here until our 1979 survey of 
the Housatonic River. B. McCabe's 1940 surveys did not find the species. It 
is presumably a recent introduction resulting from bait-bucket releases. The 
species appears to be common and established in the Housatonic, where 
it is now known from over 10 sites. This species was also first noticed in 
Quabbin Reservoir when the MDFW collected a single juvenile from the 
east shore in the early 1980s. Our 1989 shoreline samples of Quabbin show 
that it is now the most common minnow in the area of the reservoir that 
we surveyed. Bluntnose Minnows are also known from Little Alum Pond, 
Brimfield. 

notes. Bluntnose Minnows are important forage fish throughout their na- 
tive range. They are widely used as bait fish, but generally do not tolerate 
crowded bait-buckets. When they are propagated in ponds with artificial 
nest sites, up to 250 pounds per acre have been produced. 

references. Becker 1983, Scott and Crossmann 1973 (biology); Keast 
and Webb 1966 (feeding); Hubbs and Cooper 1936 (behavior); Dobie et al. 
1956 (propagation). 



Family and Species Accounts 121 



Fathead Minnow introduced 

Pimephales promelas Rafinesque 1 820 plate 24 




identification. Similar to Bluntnose Minnows, Fathead Minnows also 
have the well- developed but short first dorsal ray separated from the first 
principal ray (in adults) (see key Figure 8a) and small, irregular, crowded 
scales from the nape to the dorsal fin. Fathead Minnows have a chunkier 
body, and a terminal rather than slightly subterminal mouth; they lack 
the crosshatched scale outlines and the prominent basicaudal spot of 
Bluntnose Minnows. They have a dark peritoneum and a coiled gut. A 
red-orange form is marketed in pet and bait stores as an "orange tuffy." 

selected counts. D 8; A 7; Scales 9/41-54/9; PT 0,4-4,0. 

size . The Fathead Minnow generally reaches 4 inches TL, but most of the 
Massachusetts specimens that we have examined are one-half that size. 

natural history. Fathead Minnows are spring and summer spawners, 
beginning when water temperatures are above 59°F. Like Bluntnose Min- 
nows, Fathead Minnows deposit their eggs on the underside of underwater 
objects. When necessary, the males excavate a cavity under the object to 
create a nest, using their snouts and tails. The underside of the nest is 
cleaned, and finally rubbed with a dorsal pad that contains mucous cells. 
The exact function of the mucous is unknown, but it may assist in egg at- 
tachment or chemical location of the nest, or it may prevent fungus and in- 
crease egg survival. Spawning is nocturnal, and a female may deposit 100 to 
300 eggs in the nest at a time, after which she is chased away. The adhesive 
eggs stick to the roof of the nest or to other eggs. Up to 13,000 eggs in differ- 
ent stages of growth have been found in a single nest. Males actively guard 
the nest and tend the eggs by cleaning, circulating water, and removing un- 



122 Inland Fishes of Massachusetts 




Fathead Minnow. 

fertilized or dead eggs. Young hatch in four to six 
days and reach small adult size by the end of sum- 
mer. Fathead Minnows are short-lived; few live as long {;V^ 
as two years. Fathead Minnows are omnivorous, eating 
plant material, particularly algae and detritus, and smaller invertebrates. 
The proportion of each type of food varies with the season, the age of the 
fish, and the locality. 



x- 



■y. 



distribution and abundance. The Fathead Minnow is native to 
much of North America west of the Hudson Drainage. It is not native to 
Massachusetts. The species was first recorded from Massachusetts by 
P. Mugford in 1969. We did not find confirming specimens until 1979, when 
a population was found at the junction of the Green and Housatonic rivers 
in Great Barrington. They are common where they are found in the Housa- 
tonic but known from only a few sites. Reproducing populations were 
found in the Concord Drainage and in a pond on the University of Massa- 
chusetts' Amherst campus during the late 1980s. A single adult was also col- 
lected from a small tributary to the Connecticut River, Agawam, in 1980. 
These Massachusetts records are probably the result of bait-bucket releases. 

notes . Fathead Minnows are hardy and can endure relatively low oxygen 
levels, high levels of pollutants, and a wide spectrum of pH levels. They are 
intensively propagated and widely used as a bioassay organism, are an ex- 
cellent forage fish, and are a popular bait fish in the Midwest. Unlike the 
Bluntnose Minnow, the Fathead Minnow can survive crowded bait-buckets 
for many hours. It has been propagated in great quantity, even in tertiary 
sewage treatment ponds. Under these conditions, 100 pounds of stocked 
Fathead Minnows have produced up to 7,200 pounds in three months. 



Family and Species Accounts 123 



references. Becker 1983 (general); Smith and Murphy 1974 (dorsal pad); 
Andrews and Flickinger 1974 (spawning); Dobie et al. 1956 (propagation); 
Mugford 1969 (Massachusetts record). 



Blacknose Dace Native 

Rhinichthys atratulus (Hermann 1804) plate 21 




identification. Blacknose Dace have a barbel at each corner of the 
mouth, and a band of tissue (frenum) connects the upper lip to the snout. 
They are most similar to the Longnose Dace but can be distinguished by 
the relative length of the snout, the eye size, the position of the eye in rela- 
tion to the mouth (see key Figure 13b), and the pigment stripe on the snout. 
A dark stripe running around the snout, through the eyes, and along most 
of the midbody separates the olive-brown back and a silvery- white belly. In 
breeding males, the pectoral, pelvic, and anal fins are orange. 

selected counts. D 8; A 7; Scales 10-12/52-60/7-9; PT 2,4-4,2. 

size . Blacknose Dace are small, usually only reaching 3 inches TL. The 
largest one that we have examined is about 4 inches TL (87 mm SL). 

natural history. Blacknose Dace can be found in almost every hill 
stream in central and western Massachusetts. When disturbed, Blacknose 
Dace quickly disappear under rocks, boulders, or logs, only to return in a 
few minutes. Blacknose Dace begin spawning in early June when they leave 
the deeper pools and gather in and around riffles. Males guard a small terri- 
tory, and there is constant chasing. Several males may chase a single female, 
and if ready to spawn, the female slides up beside one of the males. The 
spawning act lasts one or two seconds; eggs and milt are simply broadcast 
into the water column. Nests are never built; however, Blacknose Dace are 



124 Inland Fishes of Massachusetts 




Blacknose Dace: open circles indicate known locali 
ties where species was not found during our post- 
1969 surveys; not all solid circles were resurveyed. 



f . 



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'V t # 



known to spawn over the stone nests made by Fall- ? :C 

fish. After spawning, both male and female Blacknose 
Dace settle on the bottom to rest momentarily and then quickly resume 
spawning with the same or different partners. Females contain an average 
of 750 small eggs (0.03 inches in diameter), and fry are less than 0.25 inches 
long when they hatch. Blacknose Dace feed on a wide variety of aquatic in- 
vertebrates and terrestrial insects. Aquatic fly larvae are a favored prey. 
Blacknose Dace may live for three to possibly four years. 






distribution and abundance. In Massachusetts, Blacknose Dace 
are by far the most common stream minnow, occurring from the Hudson to 
the Blackstone drainages and north through western portions of the Merri- 
mack River Drainage. In the eastern portion of the state, Blacknose Dace 
are now found only in five streams tributary to the Merrimack River, and in 
four streams in the Concord-Assabet River Drainage. Blacknose Dace are 
notably absent from all other Massachusetts coastal drainages. 

n o t e s . A number of Massachusetts fishes are sometimes infested with 
black spot disease, which is often indicative of stressed habitats. This dis- 
ease is common in Blacknose Dace; individuals are sometimes almost en- 
tirely covered with the diagnostic small black spots, especially when they 
are trapped in pools as water recedes. The spots, which can be found on the 
body or the fins, are the cysts of a trematode parasite, Neascus sp. These 
parasites are passed from bird droppings, to snails, to fishes, and back to 
birds, such as the Belted Kingfisher, that prey on fishes. 



Family and Species Accounts 125 



references. Houde 1964 (black spot, MA); Raney 1940b (breeding); Reed 
and Moulton 1973 (age and growth, MA); Traver 1929 (life history). 



Longnose Dace Native 

Rhinichthys cataractae (Valenciennes 1842) plate 22 




identification. Longnose Dace have a barbel at each corner of the 
mouth, and a band of tissue (frenum) connects the upper lip to the snout. 
They are similar to Blacknose Dace but have a subterminal mouth and usu- 
ally lack the dark band around the snout and along the body. They can be 
positively identified by the length of the snout, eye size, and the position 
of the eye in relation to the mouth (see key Figure 13b). The stripes on the 
snout and the midlateral area are diffuse and are not prominent. Breeding 
males are orange-red at the base of the pectoral and pelvic fins, on the 
cheek, throat, and lips; an orange wash is sometimes present on the mid- 
lateral area and on the dorsal and anal fins. Longnose Dace lack papillae 
found on the lips of suckers. 

selected counts. D 8; A 7; Scales 11-13/61-75/8; PT 2,4-4,2. 

size. Adults are normally about 3 inches TL. The largest Massachusetts 
specimen recorded is from the Westfield River and is close to 6 inches TL 
(139mmSL). 

natural history. Longnose Dace are usually associated with steep gra- 
dient, cold-water streams, but they are sometimes found in lower-gradient, 
warm-water rivers. They are sometimes abundant, appearing in densities 
of almost one fish per square foot. Longnose Dace can live to five years and 
spend most of their adult lives on or near the bottom in turbulent water or 



126 Inland Fishes of Massachusetts 




Longnose Dace: open circles indicate known locali- 
ties where species was not found during our post- 
1969 surveys; not all solid circles were resurveyed. 









/,- 



3*; 



adjacent pools. The long, sloping nose and low pec- 
toral fins help to streamline their bodies in the current. 
Spawning, which starts in the spring, probably extends into early sum- 
mer. Although they do not build nests, each male guards a territory about 
10 inches in diameter. After spawning, eggs hatch in three to four days at 
70°F. Unlike the adults, the young live off the bottom during the early part 
of their lives. Their diet consists primarily of immature aquatic insects that 
cling to rocks and boulders. Longnose Dace are one of the chief predators 
of larval blackflies and midges, but they will also prey on other small 
aquatic invertebrates. 



distribution and abundance. In western Massachusetts, Longnose 
Dace are common in clear streams with riffles, boulders, and gravel, but 
have also been sampled in large numbers from lower- gradient, main stem 
rivers, including the Housatonic River, Stockbridge. Longnose Dace are ab- 
sent from almost all of the eastern part of the state except in upland tribu- 
taries to the Nashua River. They are rare in the lower Merrimack Drainage, 
where there are only two records: one from Lawrence in 1859 and one from 
Andover in 1987. The Longnose Dace may have been more common along 
the Merrimack before industrial pollution and dams. With the exception of 
one undocumented fisheries record from the upper Taunton drainage, they 
are absent from all other Massachusetts coastal drainages. 

notes. Longnose Dace were originally described from Niagara Falls and 
were given the appropriate specific name cataractae. 



Family and Species Accounts 127 



references. Cooper 1980 (development); Reed 1959 (diet); Reed and 
Moulton 1973 (age and growth); Scott and Crossman 1973, Jenkins and 
Burkhead 1993 (general). 



Rudd 

Scardinius erythrophthalmus (Linnaeus 1758) 



Introduced 
plate 8 




identification. Rudd are deep-bodied, compressed fish with a down- 
curved lateral line. They closely resemble Golden Shiners, but Rudd lack 
the fleshy, scaleless area on the ventral midline between the pelvic fins and 
the anus. Adults are silvery with red fins. Golden Shiners also have smaller 
scales, more gill rakers, and only one row of pharyngeal teeth (see key Fig- 
ure 4b). 

selected counts. D 9-1 1; A 11-14; Scales 7-9/38-41/3-5; GR 10-13; 
PT 3,5-5,3. 

size. Rudd commonly grow to 10 or 12 inches TL but may reach 18 inches 
TL. 

natural history. Almost nothing is known about the natural history of 
the introduced Rudd in North America. In Eurasia, it inhabits lakes, ponds, 
and slow- flowing waters. Rudd spawn in the spring over submerged vegeta- 
tion, usually between April and June. Adults do not mature until they are 
three to four years old or over 5 inches TL. They are principally midwater 
and surface fish, feeding mostly on insects, crustaceans, and filamentous 
algae. Rudd have been documented as hybridizing with the native Golden 
Shiner in North America. 



128 Inland Fishes of Massachusetts 




Rudd. 



DISTRIBUTION AND ABUNDANCE. Rudd are 

native to areas of temperate Eurasia and were his- 
torically introduced to New York and Maine. By the 
mid-1980s, Rudd were found in some 14 US states, due to 
bait fish releases. It is unknown if they are reproducing in all of these states. 
The Rudd was imported to Massachusetts as a bait minnow at least as early 
as the late 1980s and confirmed in the wild by our surveys of the Charles 
River in Cambridge in 1991, when two specimens were collected: an adult 
(206 mm SL) and a young (88 mm SL) found on different dates. The presence 
of both juveniles and adults over several years confirms reproduction in the 
lower Charles River. A record exists of the species from Benton Lake, Otis. 






notes. Because this minnow was used for bait for a number of years be- 
fore its importation was prohibited by MDFW in 1990, there is a good possi- 
bility that it will be found reproducing in other areas of the state. 

references. Courtenay and Stauffer 1984 (US introductions); Burkhead 
and Williams 1991 (hybrids, identification); Hartel 1992 (Massachusetts 
records); Wheeler 1969 (biology). 



Family and Species Accounts 129 



Creek Chub Native 

Semotilus atromaculatus (Mitchill 1818) plate 19 








identification. Creek Chub are similar to Fallflsh, which also have 
a leaflike fleshy barbel in the groove behind the upper jaw (see key Fig- 
ure 1 lb). Care must be taken in looking for the barbels; they may rarely be 
absent from either side or both sides. Creek Chub have a distinctive, small 
spot near the anterior base of the dorsal fin and more than 50 lateral line 
scales. Young Creek Chub have a lateral band from the snout to the caudal 
base that often ends in a basi- caudal spot. Breeding males darken dorsally 
and have a yellow to rosy wash laterally on the body. 

selected counts. D 8; A 8; Scales 8/50-62/5; PT 2,5-4,2. 

size . Adult Massachusetts Creek Chub are usually 4 to 5 inches TL. The 
largest Massachusetts specimen that we have seen measured about 
6.5 inches TL (136 mm SL). However, they are known to grow as large as 
12 inches TL in other parts of their range. 

natural history. In Massachusetts, Creek Chub are most often found 
in small streams with gravel bottoms. Spawning takes place in the spring 
when water temperatures range from 54° to 61°F. Nest building has been 
described by Ross (1977a:36) as follows: "...male Creek Chubs move onto 
gravel runs" and the male "...forms a nest depression by carrying sand and 
gravel from the nest in his mouth. Once a depression is formed, the male 
removes gravel only from the downstream edge of the depression. This ma- 
terial is deposited on the upstream edge of the depression in the area where 
spawning occurs. Thus, eggs that are released in the nest are subsequently 
covered with a layer of gravel which is moved by the male." Creek Chub oc- 



130 Inland Fishes of Massachusetts 









V*W? 



'•*< 



...... 




• X)P>^t^ 4/ v K^V.., v l / ' aM^~- vk^' r ' 






Creek Chub: open circles indicate known localities 
where species was not found during our post- 1969 
surveys; not all solid circles were resurveyed. 



* V / v.. > !)\]f 

\ ) in 



/,-' 



-•;>••::■-:•- 



•<Cv. 



- 



casionally respond with aggressive displays to other ■; 
fish species entering their nest area, especially near the 
time of spawning. However, they always respond to other male Creek 
Chub. Creek Chub are opportunistic, feeding at all depths in the stream, 
most intensively in the evening. Their diet includes a wide range of aquatic 
insect larvae and pupae, fishes, and mollusks. Burrowing bottom organisms 
are taken to a lesser extent because the Creek Chub seems to rely on sight 
to find food. By their third summer, Creek Chub average about 5 inches, 
and they probably live longer than four years. 



/-' 



distribution and abundance. In Massachusetts, Creek Chub are 
found in most major river drainages west of the Connecticut River. East of 
the Connecticut River, Creek Chub are much less common. Only four re- 
cent records exist from the Chicopee River Drainage and two records from 
the Millers River Drainage. 

references. Barber and Minckley 1971 (food); Dinsmore 1962 (life his- 
tory); Johnston and Ramsey 1990 (relationships); Maurakis et al. 1990 
(nests); Reighard 1910, Ross 1977a, 1977b (behavior). 



Family and Species Accounts 131 



Fa 1 1 fish 

Semotilus corporalis (Mitchill 1817) 



Native 
plate 18 




identification. Fallfish are most similar to Creek Chub, which also 
have a leaflike, fleshy barbel in the groove behind the upper jaw (see key 
Figure lib). The barbels have to be looked for carefully because they may 
be rarely absent from either or both sides. Opening the mouth and direct- 
ing a small jet of air into the groove behind the posterior area of the maxilla 
will make the barbel easier to observe. Adult Fallfish have diagnostic dark 
marks at the base of each scale (see key Figure 14a) and fewer than 50 lateral- 
line scales. Fallfish are silvery with a dark olive-brown to almost black dorsal 
area. Young have a pronounced lateral band. 

selected counts. D 8; A 8; Scales 7/43-50/5; PT usually 2,5-4,2 

size . This species is Massachusetts' largest native minnow. Adults just un- 
der 1 foot long are common. The largest recorded Massachusetts specimen, 
from Quabbin Reservoir, measured 19 inches (462 mm SL). 

natural history. In Massachusetts, Fallfish are most often found in 
rivers and steams with rock and gravel substrates, but some populations 
occur in larger ponds and reservoirs. Adults migrate to areas with rock and 
gravel substrate in the spring. Dominant males begin building nests in mid- 
April, when water temperatures are above 59°F. Males dig a pit, then pick 
up stones weighing up to 6 ounces in their mouths and drop them in the 
pit to form the nests. The nests may be as small as 1 foot to over 4 feet in 
diameter and almost 2 feet high. The shape of the nest depends on its loca- 
tion; nests built in current are often elongated downstream by the force of 
the water, while those in quiet water are dome shaped. Fallfish spawning 
involves a social hierarchy established by the behavior of the male. The 



132 Inland Fishes of Massachusetts 



'<? 






. : O^ 



Fallfish: open circles indicate known localities where (</ j "\ ./ '\J % \J I L/( 
species was not found during our post- 1969 surveys; 4^ * J f 1 

not all solid circles were resurveyed. ? V / ; 



.i\ >, ,<-~"V 



;P 



dominant male makes aggressive displays and < : X 

chases subordinate males. Female Fallfish and other c \-? 

species are not chased. The visual cue of a male carrying stones and 
dropping them into the nest triggers the female to rush onto the nest to 
spawn. Spawning is often communal with varying numbers of both sexes 
involved. During communal spawning the fishes "...form a layered aggre- 
gation, all facing upstream and in close contact with each other... with the 
dominant male. ..always located centrally, next to the nest" (Ross and Reed, 
1978:218). Fertilized eggs are adhesive, stick to the nest, and hatch in about 
six days. Fallfish are omnivorous, eating plankton until they are about 
1.5 inches TL and gradually switching to larger foods, such as algae, insects, 
crayfish, and fishes. In a study in the Mill River, Amherst, Fallfish were 
found to eat more aquatic insects during the spring and gradually switched 
to heavy feeding on terrestrial insects by midsummer. It takes five years for 
a Fallfish to reach about 8 inches TL and almost 10 years to reach maximum 
size. Males become mature in their third spring and females in their fourth. 

distribution and abundance. In Massachusetts, Fallfish are com- 
mon in the Connecticut Basin but rare in the eastern part of the state. The 
latest Charles River records date back more than 30 years. Storer (1867) 
stated that they were found in many rivers; his 1839 description is based on 
a 14-inch specimen from Walpole (Charles River Drainage). Recent records 
of Fallfish from Cape Cod are lacking; however, two mounted specimens 
(12 and 16.5 inches TL) in the Springfield Museum of Natural History are 
labeled from "ponds on Cape Cod, 1911." 



Family and Species Accounts 133 



notes. Other fishes are attracted by the nest-building behavior of the 
Fallfish, and some actually spawn over their nests. In Massachusetts, Com- 
mon Shiners and Blacknose Dace sometimes spawn in this manner and 
may gather around Fallfish nests by the hundreds. 

references. Johnston and Ramsey 1 990 (relationships) ; Maurakis et al. 
1990 (nests); Reed 1971 (growth, development, diet); Ross and Reed 1978 
(reproduction); Ross 1983 (behavior); Smith 1985 (general). 



134 Inland Fishes of Massachusetts 



Sucker Family 

Catostomidae 



The suckers, with some 70 species, are closely related to the true minnows. 
Most suckers are endemic to North America, but two species, one in China 
and one in northeastern Siberia, are found in northeastern Asia. Most sucker 
species require relatively clean silt-free water, and at least 20 species are 
currently listed as threatened or rare in North America, in part due to habi- 
tat change. The Harelip Sucker, Moxostoma lacerum, was one of the first 
North American fishes to become extinct. It was formerly common from 
Ohio to Georgia and Arkansas, but none has been found since 1895. As the 
common name indicates, most members of this family have distinctive, 
subterminal, suckerlike mouths. The toothless lips are fleshy and can pro- 
trude into a short tube. Sucker lips are pleated with lines and bumps that 
contain tactile and other sensory organs. All suckers have comblike pha- 
ryngeal jaws with 20 or more teeth. The anal fin is set much farther back 
on the body than on most minnows. 

Reproduction occurs in spring. During this time, the adults congregate 
in spawning areas, usually upstream from the deeper pools and channels 
where they normally live or in the shallows of lakes and reservoirs. Spawn- 
ing occurs in groups of three, usually one female tightly flanked by two 
males. Fertilized eggs sink to the bottom and are buried by the movements 
of the anal and caudal fins of the adults, but nests are not built. During the 
spawning period, male suckers develop breeding tubercles, especially on 
their heads and anal fins. One specimen of a Northern Hog Sucker, Hypen- 
telium nigricans, was found in a tributary to the Connecticut River, Hadley, 
in 1953, but this introduced species has not been found since. 

references. Bruner 1991 (bibliography); Page and Johnson 1990 (spawn- 
ing behavior); Williams et al. 1989, Miller et al. 1989 (rare and extinct spe- 
cies); Smith 1992, Fink and Fink 1996 (systematics and relationships); Jenk- 
ins and Burkhead 1993 (general). 



Family and Species Accounts 135 



Key to Massachusetts Suckers 



la. Lateral line absent; scales large, less 
than 45 in midlateral series; young with 
dark dorso-lateral stripes. Creek Chub- 
sucker, Erimyzon oblongus, page 141, 
Plate 25. 




lb. Lateral line present; scales small, 
more than 55 lateral scales; young with- 
out dark horizontal stripes (not illus- 
trated). Go to 2. 




2a. Snout long, extending well ahead of 
the mouth; more than 85 scales in lateral 
series. Longnose Sucker, Catostomus 
catostomus, page 137, Plates 27, 28. 





>fii§i§ite« M ,:^. . 




2b. Snout short, barely extending ahead 
of the mouth; scales fewer than 75 in lat- 
eral-line series. White Sucker, Catosto- 
mus commersoni, page 139, Plate 26, 28. 




136 Inland Fishes of Massachusetts 



Longnose Sucker 

Catostomus catostomus (Forster 1773) 



Native, State Special Concern 
plates 27, 28 





\\' < .■' ... : 




identification. Longnose Suckers closely resemble White Suckers, but 
their longer and more pointed snout extends well beyond the subterminal 
mouth when viewed from below (see key Figure 2a). In addition, they have 
smaller scales with more than 85 in the lateral series and over 15 scales 
above the lateral line. Males have a coppery red midlateral stripe during 
breeding. 

selected counts. D 9-11; A 7-9; Scales 88-115. 

size. Longnose Suckers are usually 12 to 15 inches TL. Outside of New En- 
gland, occasional specimens may reach 2 feet TL. Adults from populations 
in small hill streams are often smaller. 

natural history. In Massachusetts, Longnose Suckers are most often 
found in the cold, clear streams of the western part of the state. Here they 
spawn over gravel in early spring. Gravid females and tuberculate males 
have been noted in the Hoosic and Deerfield rivers between May and early 
June. Adhesive eggs are deposited over the substrate, but no nest is built. 
The eggs hatch in 8 to 10 days, and the young move into midwater to feed 
on plankton. Like other species of suckers, the larvae and postlarvae have a 
terminal mouth that changes to subterminal as the fishes grow and move to 
the bottom. Based on Wisconsin data, Longnose Suckers are relatively slow- 
growing; they take four to five years to reach 8 to 10 inches TL. They may 
live well over 10 years. Longnose Suckers vacuum a wide variety of aquatic 
invertebrates and algae off the bottom, including amphipods, copepods, 
and the larvae of blackflies, beetles, mayflies, dragonflies, stoneflies, and 
caddisflies. 



Family and Species Accounts 137 




Longnose Sucker: open circles indicate known lo- 
calities where species was not found during our post- ^ i 
1969 surveys; not all solid circles were resurveyed. J y 



/,-• 



,«N 



DISTRIBUTION AND ABUNDANCE. In MaSSacllU- 

setts, the Longnose Sucker is limited to the western third 
of the state. It is fairly common in clean, cold portions of the Deerfield, 
Housatonic, and Hoosic drainages. However, records from 1940 through 
1956 show that it occurred historically in the Connecticut and Westfield 
rivers and at the mouth of the Chicopee River, where specimens have not 
been collected in recent years. The Longnose Sucker is currently listed as 
a State Species of Special Concern because of its decline in the lower Con- 
necticut Basin (Connecticut main stem, Westfield, and lower Chicopee riv- 
ers) and in parts of the Hoosic and Housatonic drainages as a result of the 
poor water quality. The pollution and habitat alteration along the main 
stems have limited surviving populations to the cleaner tributaries (D.G. 
Smith 1990, pers. comm.). 






'•- 



notes. B. McCabe recognized two subspecies of the Longnose Sucker 
from Massachusetts. C. c. catostomus was recognized as a larger form with 
smaller scales (more than 100) and C. c. nannomyzon as a smaller, dwarf 
form with larger scales (85 to 100). Although the New England populations 
have not been studied in depth, it is probable that the subspecific status is 
not warranted. 

references. McCabe 1942, 1943 (distribution, subspecies); Sayighand 
Morin 1986 (diet); Becker 1983 (general). 



138 Inland Fishes of Massachusetts 



White Sucker Native 

Catostomus commersoni (Lacepede 1803) plates 26, 28 





, "",-VTf J ' ' - 




identification. White Suckers are similar to Longnose Suckers except 
that their snouts are rounded and barely project beyond the upper lip when 
viewed from below (see key Figure 2b). White Suckers have fewer than 
75 scales in a lateral series and fewer than 11 scales above the lateral line. 
Three or more irregular lateral blotches are usually present in juveniles and 
some adults. 

selected counts. D 10-13; A6-8; Scales 53-80. 

size . Large specimens may reach 28 to 30 inches TL, but most individuals 
are less than 2 feet TL. 

natural history. White Suckers live in a wide variety of habitats in 
Massachusetts. They are most often found in ponds, lakes, and rivers, espe- 
cially if there are tributaries with gravel runs in which to spawn. Spawning 
takes place in mid-April to May in Massachusetts, when adults move up- 
stream into tributaries or into shoal areas if tributaries are not available. 
Mating has been described as a "...tremoring trio, a female tightly flanked 
on each side by a male" (Jenkins and Burkhead 1993: 641). Sexually mature 
White Suckers may not spawn every year and exhibit a wide range in size 
at maturity. Depending on size, females may carry between 20,000 and 
139,000 eggs. Young-of-the-year grow quickly and may reach 4.5 inches in 
length by the end of their first summer. Adults live up to 10 years. Food is 
mainly benthic invertebrates and fish eggs, but larval midges make up a 
portion of their diet. While detritus is also often taken in quantity, suckers 
have the ability to detect food types with taste buds and sort out and expel 
unwanted items. 



Family and Species Accounts 139 



i ft A ^^-^Trnfi^'d^^^^ 





White Sucker: open circles indicate known localities 
where species was not found during our post-1969 
surveys; not all solid circles were resurveyed. 



DISTRIBUTION AND ABUNDANCE. In MaSSachu- ? '" £ 

setts, White Suckers are found in virtually every drainage 

with the exception of Martha's Vineyard and Nantucket and several of 

the smaller mainland coastal streams. This species is abundant in many 

locations. 






notes. White Suckers are sometimes so common that they are considered 
"trash" fish. In fact, suckers are a valuable component of our aquatic eco- 
system because they reproduce in great numbers and form a large part of 
the total fish biomass in many areas. In addition, the concept of trash fish is 
erroneous since it is now generally accepted that every species has a valued 
place in the ecosystem. Large suckers also put up a good fight when hooked 
on a light rod, and their flesh, though bony, is quite good, especially in the 
spring. 

references. Bruner 1991 (bibliography); Beamish 1973 (age and growth); 
Quinn 1982 (age, spawning); Quinn and Ross 1985 (non-annual spawning); 
Jenkins and Burkhead 1993 (general). 



140 Inland Fishes of Massachusetts 



Creek Chubsucker Native 

Erimyzon oblongus (Mitchill 1814) plate 25 



/;•>■; 



identification. Creek Chubsuckers are superficially similar to min- 
nows but have pleated and fleshy lips, a posteriorly placed anal fin, and a 
higher number of dorsal fin rays. The mouth is almost terminal, the scales 
are large, and lateral-line pores are lacking. Young have a dark brown mid- 
lateral stripe from the snout to the base of the caudal fin and a second, less- 
defined stripe between it and the dorsal midline on a golden-bronze to 
yellow-brown background. Adults lose these colors although occasional 
diffuse vertical blotches may be present along the sides of the body. 

selected counts. D 1 1-13; A 7; Scales 41-44 (midbody scales unpored). 

size . Creek Chubsuckers are generally less than 9 inches total length, but 
individuals as large as 18 inches are occasionally encountered outside of 
Massachusetts. The largest Massachusetts specimen that we examined, 
from Federal Pond in Carver, measured 14 inches TL (283 mm SL). 

natural history. Creek Chubsuckers are typically found in creeks, 
streams, and lakes with moderate quantities of aquatic vegetation but are 
also found in the clear waters of lakes and reservoirs. Creek Chubsuckers 
feed on plant material and a variety of aquatic and terrestrial invertebrates. 
As their almost terminal mouth might suggest, they spend a considerable 
amount of time feeding above the bottom. Creek Chubsuckers spawn in the 
early spring, but we have never observed them spawning in Massachusetts. 
From other studies, we know that males grow as large as females, develop 
large breeding tubercles on the head, and usually spawn in pairs. Creek 
Chubsuckers do not build nests but defend territories over gravel runs. In- 



Family and Species Accounts 141 




¥' s M$m*^. 



A. 



it} HVf* 



Creek Chubsucker: open circles indicate known lo- 
calities where species was not found during our post- 
1969 surveys; not all solid circles were resurveyed. 



) } 



)y 






; .- - -/ 



rv • 



dividual females have been found to carry up to 

72,000 eggs. No parental care is given the eggs or young, 

and after hatching, the young form schools, feed on zooplankton, and 

gradually move downstream. After spawning, the adults return to the 

downstream habitats where they live the rest of the year. Female Creek 

Chubsuckers can live at least seven years. 



: . ~X.y 



distribution and abundance. In Massachusetts, this species is 
relatively more common east of Quabbin Reservoir but is not known from 
Cape Cod and the Islands. Surveys between 1970 and 1991 have failed to 
find this species at a number of localities in Massachusetts where they were 
found prior to 1969. The areas where they were not collected are scattered 
throughout their local range, and their absence cannot be attributed to any 
particular environmental factors. However, they are known to be sensitive 
to pollutants, especially silt. 

remarks. Historical reports listed the Lake Chubsucker, Erimyzon sucetta, 
found in the Great Lakes drainages and along the southern East Coast, as 
part of the Massachusetts fauna. Our review of a large number of speci- 
mens demonstrates that the Lake Chubsucker is not found in Massachu- 
setts and that the early accounts were simply mistaken identifications of 
Creek Chubsuckers. 

references. Bruner 1991 (bibliography); Gilbert and Wall 1985 (status, 
southeastern US); Hubbs 1928, (systematics); Page and Johnson 1990 (re- 
production); Wagner and Cooper 1963 (population density, growth). 



142 Inland Fishes of Massachusetts 



Bullhead Catfish Family 

Ictaluridae 



Catfishes belong to the order Siluriformes and are closely related to the 
minnows and suckers. They are a large group of fishes containing over 
30 families and at least 2,000 species. Most catfishes, except three families, 
inhabit freshwaters in temperate to tropical regions of the world. Catfishes 
have one to four pairs of barbels and heavy skull bones; they typically have 
sharp spines in their dorsal and pectoral fins and usually lack scales. Catfish 
spines are often serrated or barbed and have poison glands that can inflict 
a painful sting. Only two native catfish families are known from New En- 
gland: the Ictaluridae, endemic to North America, and the Ariidae, a world- 
wide group of marine catfishes. The sea catfishes, found in coastal waters 
south of Cape Cod, can be identified by their lack of nasal barbels. The ex- 
otic Asian Walking Catfish, Clarias batrachus, has also been caught by local 
anglers but is not reproducing in Massachusetts. The presence of these air- 
breathing catfishes in Massachusetts is the result of the release of aquarium 
specimens. Local ictalurids are largely nocturnal and rely heavily on their 
barbels as sensory devices. The barbels, supported by a small cartilaginous 
rod and moved by small muscles, have special cells to detect tactile and 
chemical signals from the environment. These cells, which are also found 
on the body, play an important role in orientation, schooling, breeding, 
electrolocation, and feeding. The barbels, sometimes called "whiskers," 
cannot sting. 

references. Alexander 1965 (morphology); Raney 1957 (NY); Jones et al. 
1978 (early life history, development); Lundberg 1970, 1975, 1982, Fink and 
Fink 1996 (systematics and relationships); Kendall 1910 (habits, culture, 
commercial importance); Langlois 1936 (growth); Birkhead 1972 (toxic 
spines). 



Family and Species Accounts 143 



Key to Massachusetts Catflshes 

Note: One of the key characteristics separating catfish species is the total number of anal fin 
rays, including those that may he hidden beneath the skin at the anterior end of the anal 
fin. Identification of large catfishes can be difficult due to worn and broken tail fins, barbels, 
or pectoral spines, which are used for identification. Inspection of bony processes of the 
skull or pectoral girdle may be necessary to identify these questionable specimens, and dis- 
section or radiography may be necessary when a specialist examines the specimens. 



la. Adipose fin a flag-like fleshy lobe, 
well separated from caudal fin; tail 
squared, rounded, or forked; adults to 
over 24 inches TL. Go to 2. 

lb. Adipose fin long, low, and "keel-like," 
nearly continuous with caudal fin; tail 
squared or rounded; adults small, seldom 
over 6 inches TL. Madtoms Noturus. 
Go to 6. 



1a 



1b 



2a. Tail deeply forked, lobes pointed; 
anal fin with 24 to 30 rays; bony ridge 
connecting skull and origin of dorsal fin; 
head relatively small and narrow; young 
with small spots; larger adults blue-black 
in color without spots. Channel Catfish, 
Ictalurus punctatus, page 151, Plate 34. 




2b. Tail at most moderately forked, lobes 
more or less rounded; anal fin usually 
with less than 25 rays; area in front of 
dorsal fin compressible, without con- 
necting bony ridge; head large and 
broad; sometimes mottled but never with 
small spots. Go to 3. 



3a. Tail moderately forked, upper lobe 
usually longer and rounded; gill rakers 
18 to 23; head wide and massive; chin 
barbels light colored. White Catfish, 
Ameiurus catus, page 146, Plate 31. 

3b. Tail only slightly indented, square or 
rounded; gill rakers usually fewer than 19 
(except the Black Bullhead, Ameiurus 
melas); head large but never massive; 
chin barbels light or dark. Go to 4. 





144 Inland Fishes of Massachusetts 



4a. Chin barbels whitish; rear edge of 
caudal fin nearly straight or slightly 
rounded; anal fin rays 24 to 28. Yellow 
Bullhead, Ameiurus natalis, page 148, 
Plate 33. 

4b. Chin barbels dark; rear margin of 
caudal fin slightly notched and squarish; 
anal fin rays fewer than 25. Go to 5. 




5a. Well-developed serrations on poste- 
rior edge of pectoral spine; gill rakers 
13 to 15; lacking dark pigment on anal fin 
membrane. Brown Bullhead, Ameiurus 
nebulosus, page 149, Plate 32. 



5b. Serrations on pectoral spine poorly 
developed or absent; gill rakers 16 to 21; 
dark pigment on membranes of anal fin. 
Black Bullhead, Ameiurus melas. See 
comments under Brown Bullhead, 
page 149. 




6a. Body short and stout with large head; 
tail oval and paddlelike, broadly joined to 
adipose fin; vertical fins without dark 
edges. Tadpole Madtom, Noturus gyri- 
nus, page 153, Plate 35. 

6b. Body more slender and elongate; tail 
square, only narrowly joined to adipose 
fin by a low keel; vertical fins often with 
dark margins. Margined Madtom, Notu- 
rus insignis, page 155. 




Family and Species Accounts 145 



White Catfish Introduced 

Ameiurus catus (Linnaeus 1758) plate 31 



identification. The White Catfish has a large, wide head with whitish 
chin barbels. The tail is moderately forked with rounded lobes. The upper 
lobe is often slightly longer than the lower. The anal fin is relatively short 
and rounded (less than 25 rays), and this species has 18 to 23 gill rakers. 
Teeth on pectoral spines are often hooked in young. 

selected counts. D i,6; A 22-24; GR 18-23. 

size. White Catfishes are intermediate in size between Channel Catfishes 
and bullheads but have more massive heads. Adults are usually 10 to 
18 inches TL and weigh up to 4 pounds. The Massachusetts angling record 
for a White Catfish is one that weighed 9 pounds, 3 ounces; it was taken at 
Baddacook Pond, Groton, in 1987. The second largest specimen, taken in 
1988, weighed 7 pounds, 11 ounces. 

natural history. The White Catfish inhabits waters that are intermedi- 
ate between those preferred by Channel Catfishes and bullheads. In Massa- 
chusetts, they are found in the main stems of moderately large rivers and a 
few large ponds. They most frequently inhabit areas with slower currents. 
They are not found in large numbers either within dense beds of vegetation 
or in small, muddy, shallow ponds. White Catfishes can tolerate low levels 
of salinity and may occupy slightly brackish coastal streams and estuaries. 
Spawning occurs in early summer as water temperatures approach 70°F. 
Large nests are built near sand or gravel banks. Like other catfishes, they 
will also spawn in discarded containers and natural cavities. The eggs hatch 
in six to seven days. The White Catfish is omnivorous, with younger individ- 
uals feeding on aquatic invertebrates, plants, and fish eggs. Adults often 



146 Inland Fishes of Massachusetts 



^ : L- 




' \ '/ i V ' : v \\ <• ! %' ■■■ ' 

■~<. II.. ' : V \ \ S ~: \YNJS V ' 7"*»~*VV '' 









White Catfish: closed circles indicate verified records; 

open circles indicate unverified fisheries reports. V// ; «i / H 

prey on small fishes; adult smelt were found in the ^t ? 
stomach of a large White Catfish taken in the Charles 
River, Cambridge. Sexual maturity may be reached at 7 to 
8 inches TL. 







11 

y " 


/,' 


/•" " : - 




.,< 


~;'%y. *#L 


""' ! ( 






distribution and ab undance . White Catfishes were introduced as a 
sport fish in Massachusetts between 1910 and 1949. Reproducing popula- 
tions currently inhabit the Connecticut, Merrimack, Blackstone, and Charles 
rivers. In addition, there are records from a number of ponds: Baddacook 
Pond, Groton; Whitehall Reservoir and North Pond, Hopkinton; Quaboag 
Pond, Brookfield; and Mashpee-Wakeby Pond on Cape Cod. A number of 
these sites are based on fisheries or sportfishing records, and the specimens 
have not been retained or critically examined. 

notes. White Catfishes are active year-round, and they are often taken 
while people are icefishing on some of the large ponds. 

references. Sprenger 1990 (Merrimack); Schwartz 1964 (salinity toler- 
ance); Schwartz and lachowski 1965 (age and growth); Miller 1966a (sum- 
mary); Kendall and Schwartz 1968 (temperature, salinity tolerance); Kellogg 
and Gift 1983 (temperature and growth relationships); Arini 1994 (Charles 
River) . 



Family and Species Accounts 147 



Yellow Bullhead Introduced 

Ameiurus natalis (Lesueur 1819) plate 33 




identification. Yellow Bullheads have square or slightly indented tails 
with rounded corners and white or light- colored chin barbels. The anal fin 
is long (24 to 28 rays), and they have a moderate number of gill rakers. 

selected counts. D i,6; A 24-28; GR 12-18. 

size. Yellow Bullheads are a relatively small species; adults reach 8 to 
12 inches TL. 

natural history. Yellow Bullheads generally inhabit moderately or 
heavily vegetated areas of low- gradient streams and shallow bays of ponds 
and lakes. They prefer clear water; however, they are somewhat tolerant of 
silly conditions, particularly in the absence of other competing bullhead 
species. Sexual maturity is attained by the third summer when they reach 
7 to 1 1 inches TL. Spawning occurs in mid-May to early June and lasts about 
two weeks. Nests are constructed at depths of 1.5 to 4 feet, and the eggs 
hatch in 5 to 10 days, depending on the water temperature. Like other bull- 
heads, Yellow Bullheads forage most actively at night. Their diet includes 
insects, crustaceans, mollusks, and small fishes, as well as some plant 
material. 

distribution. Yellow Bullheads were first introduced into Massachusetts 
waters in 1917. They are currently found in eastern portions of the Millers 
and Chicopee drainages as well as in the Thames, Blackstone, Charles, and 
Merrimack drainages. They are common to abundant and sometimes out- 
number the native Brown Bullhead. 



148 Inland Fishes of Massachusetts 



• - 




Yellow Bullhead: open circles indicate known locali- 
ties where species was not found during our post- 
1969 surveys; not all solid circles were resurveyed. 






u 






S'*A'*&jM '' 



notes. This introduced species has an east-central 

distribution unlike that of any of our native species, 

which reflects its exotic nature. It is found in a block from the eastern tribu 

taries to the Connecticut Basin to the western edges of the coastal systems 

but not in southeastern areas and Cape Cod. 



^;, 



references. Fowler 1917 (breeding habits); Fish 1932 (early life history); 
Schaffman 1955 (age and growth); Todd et al. 1967, Todd 1971 (chemical 
communication); Reynolds and Casterlin 1977 (activity cycles); Miller 
1966b, Trautman 1981 (summaries). 



Brown Bullhead 

Ameiurus nebulosus (Lesueur 1819) 



Native 
plate 32 




identification. Brown Bullheads have square or only slightly indented 
tails with rounded corners, brown to black chin barbels, well-serrated pec- 
toral spines, and 13 to 15 gill rakers. They are most similar to the Black Bull- 
head, A. melas, which has 15 to 21 gill rakers and weakly serrated pectoral 



Family and Species Accounts 149 



K. 







• 



Brown Bullhead: open circles indicate known locali- 
ties where species was not found during our post- 
1969 surveys; not all solid circles were resurveyed. 

fin spines. Serrations can be checked by running a 

fingernail along the back side of a pectoral spine. Adults 

vary from yellow-brown to almost blue-black dorsally and pale-yellow 

to white ventrally. Some are mottled brown. 




at 



••••••• i 



7** 



■ „ &\ 



selected counts. D i,6; A 18-24; GR 13-15. 

size . Brown Bullheads are medium-sized catfishes that usually reach 8 to 
14 inches TL. The Massachusetts sportfish award for the smaller catfishes 
is awarded for "bullheads," which include Brown and Yellow bullheads. 
The tied record is held by two Brown Bullheads, each weighing 3 pounds, 
8 ounces, and angled from Stiles Pond and Whitehall Reservoir in 1985 and 
1987, respectively. 

natural history. Brown Bullheads inhabit lakes, ponds, and back- 
waters of streams and rivers, with aquatic vegetation and sandy to muddy 
bottoms. Brown Bullheads are hardy fish and tolerate adverse environmen- 
tal conditions that exclude other fishes. They are able to survive water tem- 
peratures as high as 97°F in the summer as well as oxygen concentrations as 
low as 0.2 ppm during winter and reportedly are able to survive temporary 
drought conditions by burrowing into the bottom mud. Brown Bullheads 
are dormant over the winter and often bury themselves in the mud until 
spring. They spawn from late May through June when water temperatures 
rise above 65°F. Like other catfishes, a male and a female assume a head to 
tail posture during spawning. Females do not attain sexual maturity until 
they are three years old and 8 to 13 inches long. Males mature at a some- 



150 Inland Fishes of Massachusetts 



what smaller size. One or both parents guard the eggs and young; adults 
stay until the young are 1 to 2 inches TL. The young, in broods of up to 600, 
remain together in shallow water with aquatic vegetation until the end of 
the first summer. Brown Bullheads are omnivores, feeding on a wide vari- 
ety of animal and plant material, particularly during the evening hours. 

distribution. Brown Bullheads are the only catfishes native to Massa- 
chusetts. They are common to abundant and found in every major drain- 
age but are generally absent from hillstream systems. 

notes. Brown Bullheads are often called "horned pouts" by anglers in the 
Northeast. In spite of commonly heard stories, they cannot sting with the 
barbels. Some Massachusetts specimens have been, on occasion, mistak- 
enly identified as Black Bullheads, Ameiurus melas, but only one specimen 
of the Black Bullhead is confirmed from Massachusetts. 

references. Breder 1935, 1939 (reproduction); Langlois 1936 (length- 
weight); Raney and Webster 1940 (food, juvenile growth, NY); Stroud and 
Bitzer 1955, Grice 1958, Mullan 1959, McCaig et al. 1960 (harvest, manage- 
ment, competition, MA); Loeb 1964 (burrowing behavior); Keast and Webb 
1966 (feeding ecology); Hartel 1992 (Black Bullhead records). 



Channel Catfish introduced 

Ictalurus punctatus (Rafinesque 1818) plate 34 




identification. Channel Catfishes have a deeply forked caudal fin with 
pointed lobes, relatively narrow heads, long anal fins with over 24 rays, and 
dark-colored chin barbels; the barbels at the corner of the mouth are greater 
than three times as long as those near the nostrils. In addition, they can be 
distinguished from all bullheads by having a bony ridge that connects the 



Family and Species Accounts 151 




Channel Catfish: closed circles indicate verified 
records; open circles indicate unverified fisheries 
reports. 



skull to the origin of the dorsal fin. Color is variable 
with age, sex, and habitat. The body is silver-blue dor- 
sally to yellow- white ventrally, but larger fishes are often 
uniformly dark. Scattered black spots are present on the sides of the body 
in fishes smaller than 12 inches TL. 



■/f\ 



selected counts. D i,6; A 24-30; GR 13-18. 

size. Channel Catfishes are the largest members of the family in Massa- 
chusetts. Weights of 2 to 5 pounds are not uncommon, and individuals may 
reach 10 to 15 pounds. The Massachusetts sportfish record is a 26.5-pound 
Channel Catfish that was angled from the Ashfield Lake in 1989. Adults may 
be over 30 inches TL in part of their range. 

natural history. Channel Catfishes inhabit large bodies of water with 
sand or gravel bottoms that are relatively free of silt. They are seldom found 
in the shallower, more turbid, vegetated areas frequented by bullheads. 
Adults tend to be migratory and move into spawning areas in the late spring 
to early summer when water temperatures approach 70°F. Spawning takes 
place in secluded nests built by the males in holes beneath undercut banks, 
rocks, or logjams. Broken drainage tiles or large cans are also used as spawn- 
ing sites. Young Channel Catfishes reach 2 to 5 inches TL during their first 
summer and 12 to 18 inches TL before reaching maturity at four to seven 
years of age. Some individuals may live more than 15 years. 

Channel Catfishes feed throughout the water column, from the bottom to 
the surface. Though considered nocturnal feeders, Channel Catfishes also 



1 52 Inland Fishes of Massachusetts 



feed during the daytime, and they probably use their larger eyes to feed by 
sight much more than the other catfishes. Channel Catfishes feed on a wide 
variety of plant and animal matter: the young feed primarily on aquatic 
insects, while adults are omnivorous, with fish comprising a large part of 
their diet. 

distribution. In Massachusetts, Channel Catfishes were introduced 
into the Connecticut River between 1920 to 1960. Since that time, their 
range has expanded to include lower portions of major tributaries to the 
Connecticut River (Chicopee-Quaboag rivers, Deerfield River). They are 
also found in a number of larger lakes and ponds such as Baddacook Pond, 
Groton, and Quaboag Pond, Brookfield. Many records are based on infor- 
mation from fisheries or sportfishers that has not been critically examined. 

notes . The "farm-raised" commercial catfish meat purchased in most 
markets is from Channel Catfishes. As one of the largest freshwater fishes 
in the state, the Channel Catfish is actively sought for sport and food. 

references. Bailey and Harrison 1948 (food habits); Marzolf 1955 (age 
and growth); Clemens and Sneed 1957 (spawning); Moss and Scott 1961 
(oxygen requirements); Nowicki and Mann 1989 (Connecticut River, MA); 
Arini 1994 (Charles River, MA); Schwartz 1964 (salinity); Sneed 1964 (hy- 
bridization); Lewis 1976 (food). 



Tadpole Madtom introduced 

Noturus gyrinus (Mitchill 1817) plate 35 



identification. Tadpole Madtoms have well-developed procurrent 
caudal fin rays that form an oval, paddle-like tail broadly joined to the adi- 
pose fin. They are uniformly dark in color with a rather chunky body that 
somewhat resembles a tadpole of a frog or toad. 



Family and Species Accounts 153 




Tadpole Madtom. 



selected counts. D i,5-6; A 12-18; GR 5-10. 



size. Tadpole Madtoms are Massachusetts' smallest 
catfish. Most adults are less than 4 inches TL, although the 
species has been known to reach almost 5 inches TL. 



natural history. In Massachusetts, Tadpole Madtoms are most often 
found in ponds and well-vegetated sections of slow- flowing streams. How- 
ever, they also inhabit sandy bottomed streams with sparse plant life. Re- 
production has not been studied in Massachusetts, but in other parts of 
their range, this species spawns in the late spring and early summer. Tad- 
pole Madtoms deposit a cluster of relatively large eggs in nest cavities. The 
adults and egg masses containing up to 100 eggs are often found inside dis- 
carded tin cans. Both sexes are known to guard the nest. Tadpole Madtoms 
are nocturnal and usually feed on a range of small invertebrates, especially 
isopods and larval midges. Like many small species, Tadpole Madtoms 
have a short life span, lasting only two to three years. 

distribution and abundance. This species was first found in Massa- 
chusetts in 1939 at Howe Pond, Spencer, near the headwaters of the Chico- 
pee Drainage. Since then, madtoms have been found as far downstream as 
the Red Bridge Dam on the Chicopee main stem, and in a number of locali- 
ties in the French River (Thames Basin). Tadpole Madtoms are common in 
local areas of both drainages, but they have never been reported in any other 
parts of the Connecticut Basin in either Massachusetts or Connecticut, nor 
have they been found in the Thames Drainage below the dam on the French 
River near the Massachusetts state line. The thousands of unidentified small 
"horned pouts," stocked in the 1930s, are probably the source of the intro- 



1 54 Inland Fishes of Massachusetts 



duction. Howe Pond, itself, received 5,650 catfishes. New Hampshire's pop- 
ulation was considered introduced when first reported in 1938. Based on 
these facts, we consider the Tadpole Madtom an introduced species in Mas- 
sachusetts. However, it is also possible that madtoms found in New England 
are disjunct and relict to populations on the southern Atlantic Coastal Plain 
(Schmidt 1986). 

notes. Like some other catfishes, the madtoms have sharp pectoral spines 
associated with venom glands. Tadpole Madtoms are especially unpleasant 
if handled carelessly. Even a 2- or 3-inch specimen can produce a painful 
sting, like that of a wasp, which can last for 15 or more minutes. 

references. McCabe 1948 (first MA record); Bailey 1938 (NH records); 
Schmidt 1986 (zoogeography); Birkhead 1972 (toxic spines); Smith 1985 
(general, NY); Taylor 1969 (systematics and description). 



Margined Madtom introduced 

Noturus insignis (Richardson 1836) 






identification. Margined Madtoms are slender and elongate, have a 
square tail fin, and have small procurrent rays that join the tail to the adi- 
pose fin by a low keel. They also have an overshot upper jaw and barbs on 
the posterior edges of the pectoral spines. Margined Madtoms are slate- 
gray to yellow-tan dorsally with lighter sides shading to creamy- white be- 
neath the head and belly. The pectoral, dorsal, anal, and caudal fins are of- 
ten outlined or margined with black. 

selected counts. D i,5-6; A 15-21; GR 6-10. 

size . Margined Madtoms generally grow to 5 inches TL and rarely exceed 
6 inches. 



Family and Species Accounts 155 







Margined Madtom. 

natural history. Over most of their range, 
Margined Madtoms inhabit moderate to low cur- 
rents of larger, clearwater streams with rocks, boulders, 
or coarse gravel. Maturity is not reached until their second b 
year of life, when they are about 5 inches TL. In Pennsylvania, Margined 
Madtoms spawn in June, and, over their range, they typically nest under 
flat stones. Each clutch contains from 50 to 200 eggs. Margined Madtoms 
are active nocturnal omnivores that eat a large variety of aquatic insects 
and other invertebrates. During the daytime, they may be found lying pas- 
sively concealed beneath stones and bottom debris. Margined Madtoms live 
about four years. 



<:/' \ 



distribution and abundance. Margined Madtoms were first found 
in Massachusetts during a Division of Fisheries and Wildlife survey in late 
July 1988 when two specimens 85 and 95mm SL were collected in Crooked 
Springs Brook, Chelmsford, a tributary to the Merrimack River. R.M. Bailey 
(1938) considered the Margined Madtom to be an introduced species when 
he first found it in the Merrimack system in New Hampshire. Although it is 
possible that the New Hampshire and Massachusetts populations are dis- 
junct relicts (Schmidt 1986), we consider this an introduced species in Mas- 
sachusetts. See comments under Tadpole Madtom. 

notes. The sharp pectoral spines have a venom gland at their base, and a 
painful beelike sting can be inflicted if the fish are handled carelessly. 

references. Reed 1907, 1924 (poison spines); Fowler 1917 (breeding 
habits); Bowman 1932, 1936 (ecology, notes); Clugston and Cooper 1960 
(growth, PA); Flemer and Woolcott 1966 (food habits); Halliwell 1988 (Mer- 
rimack Drainage) . 



1 56 Inland Fishes of Massachusetts 



Pike and Pickerel Family 

Esocidae 



Pike and pickerels are most closely related to the mudminnows and dis- 
tantly related to salmonids. The pike family is small with only a single ge- 
nus, Esox, and five species that are found only in the Northern Hemisphere. 
Members of this family have elongated bodies, with the dorsal and anal fins 
set far back on the body and opposite each other. Their small pectoral and 
pelvic fins are inserted low on the body. All pickerels and pike lack an adi- 
pose fin. Their heads are long with a broad, rather flat snout, and they have 
a large mouth with numerous, well-developed teeth. The pattern of scales 
on the cheeks and gill covers is one of the primary taxonomic characters 
distinguishing species. 

All members of this family are predators; they are "wait and ambush" 
hunters that hover quietly and then dart forward with a flick of the tail. The 
larger species are top predators in their food chain and grow to impressive 
sizes. Spawning occurs in shallow areas with abundant emergent or sea- 
sonally flooded vegetation during early spring. Adhesive eggs are broadcast 
over vegetation, and no parental care is given. One of the most important 
factors in successful esocid reproduction is water level stability during 
spawning and early growth because either eggs or young may be stranded 
by abnormal water level fluctuations in the shallow water breeding areas. 

references. Wich and Mullan 1958 (life history, ecology, MA); Crossman 
and Buss 1965 (hybrids); Crossman 1978, Lundberg 1982 (relationships); 
Casselman et al. 1986 (identification, hybrids). 



Family and Species Accounts 157 



Key to Massachusetts Pike and Pickerels 

Note: Hybrids are known between all of these species, and intermediates that do not quite 
fit the key can occur (see Casselman et al. 1986 for more information). 



la. Gill covers not fully scaled; usually 10 
submandibular pores (5 on each side). 
Pike. Go to 3. 




lb. Gill covers fully scaled; usually 8 or 
fewer submandibular pores (4 on each 
side). Pickerels. Go to 2. 




2a. Snout short and convex, about equal 
to depth of head at mid eye; branchioste- 
gal rays 1 1 to 12; vertical bars on sides of 
body; teardrop below eye often slanted 
backward; lower fins red to orange. Red- 
fin Pickerel, Esox a. americanus, page 159, 
Plate 36. 




2b. Snout long and concave, always 
greater than depth of head at mid eye; 
branchiostegal rays 13 or more; adults 
with chainlike markings on sides of body; 
teardrop below eye usually vertical; lower 
fins never red or orange. Chain Pickerel, 
Esox niger, page 163, Plate 37. 




3a. Pattern of light yellow to white, bean- 
shaped spots on a darker body color. 
Northern Pike, Esox lucius, page 161. 

3b. Pattern of vertical dark bars on a sil- 
very or light body color (but pattern may 
be variable). Tiger Muskie, Esox lucius x 
E. masquinongy. See comments under 
Northern Pike, page 161. 




158 Inland Fishes of Massachusetts 



Redfin Pickerel Native 

Esox americanus americanus Gmelin 1788 plate 36 




identification. Redfin Pickerel have fully scaled cheeks and opercula; 
a short, convex snout; vertical barring on the back and sides; and reddish 
lower fins. The dark bar below the eye usually slants slightly backwards. 
Variable color is most often dark green to brown-green above, shading to 
grass-green laterally, and creamy white ventrally. Juveniles (2 inches or 
smaller) are uniformly darker than adults and do not have prominent ver- 
tical bars. 

selected counts. D 15-18; A 13-17; Scales 94-1 16; Branchiostegals 
12-13; Submandibular pores 4 (3-5). 

size. Redfin Pickerel are the smallest esocids; adults are almost always less 
than 12 inches TL. The largest Massachusetts specimen that we have mea- 
sured is just over 7 inches TL (160 mm SL). 

natural history. Prior to translocation by humans, Redfin Pickerel 
were restricted to low elevation areas with slow-moving, often naturally 
acidic streams and small ponds. Redfin Pickerel spawn early in spring when 
water temperatures approach 50°F — probably April to May in Massachu- 
setts. Spawning occurs along heavily vegetated flooded margins of small 
streams and ponds. Redfin Pickerel mature in two to three years and may 
live up to seven years. Adults prey mostly on small fishes and crayfishes; 
juveniles feed primarily on aquatic insects and other invertebrates. Vora- 
cious predators, these small pickerel use an ambush style of hunting like 
their larger relatives, but on much smaller prey. 

distribution. The Redfin Pickerel is the eastern subspecies of Esox 
americanus, which has a western subspecies called E. a. vermiculatus. In 



Family and Species Accounts 159 



*iw 




Redfin Pickerel: open circles indicate known locali- 
ties where species was not found during our post- 
1969 surveys; not all solid circles were resurveyed. 



ti* i. 



V/ 



r 



7 



uU 






- 
XQl. 



Massachusetts, Redfin Pickerel are commonly 

found throughout the coastal lowlands. There are a 

few records of this species from the floodplain of the Connecticut River, 

just north of the Connecticut state line. Redfin Pickerel found in the 

Housatonic Drainage are the result of introductions. 



\% 



% 



c 



notes. Also called "bulldog" or "mud" pickerel, this species is often mis- 
taken for juvenile Chain Pickerel. Before the introduction of Largemouth 
Bass into its Massachusetts range, Redfin Pickerel was the top predator in 
a coastal fauna that includes Swamp Darters and Banded Sunfish. Redfin 
Pickerel readily hybridize with Chain Pickerel and produce fertile hybrids, 
which are common in Massachusetts. 

references. Buss 1962, Crossman 1962 (life history, ecology); Crossman 
1966, 1978 (taxonomy and distribution); Chang 1979 (food habits); Raney 
1955 (hybrids, MA). 



160 Inland Fishes of Massachusetts 



Northern Pike introduced 

Esox lucius Linnaeus 1758 




identification. Northern Pike have scales on the cheek and on the up- 
per half of the operculum. They also have a total of 10 to 1 1 submandibular 
pores, whereas pickerels usually have 8 or fewer total pores. The color pat- 
tern of light, bean-shaped spots on a dark background separates the North- 
ern Pike from the hybrid Tiger Muskellunge ("Muskie"), which has a verti- 
cally striped pattern on the body. In young Northern Pike, the spots are less 
numerous and arranged in vertical rows that appear as bars. 

selected counts. D 15-19; A 12-15; Scales 105-148; Branchiostegals 
14-15; Submandibular pores 5 (3-6). 

size. Northern Pike are large fish, although not quite so large as Muskel- 
lunge. Lengths of 19 to 37 inches TL and weights of 2 to 12 pounds are most 
common, but some Northern Pike grow to 4 feet or more. The Massachu- 
setts angling record for Northern Pike weighed 35 pounds when caught 
through the ice in 1988 at South Pond in East Brookfield. The largest Massa- 
chusetts Tiger Muskie weighed 19.4 pounds when taken from Lake Quan- 
napowitt in 1994. 

natural history. In Massachusetts, Northern Pike generally spawn 
from late March to April, when water temperatures exceed 40°F. A large fe- 
male and one or two smaller males form a spawning group that swims into 
vegetation where eggs and sperm are released as the fish vibrate their bod- 
ies. Spawning is repeated many times over 2 to 5 days, and the fertilized 
eggs adhere to vegetation and hatch in 12 to 14 days. The half-inch-long fry 
attach themselves to vegetation for 6 to 10 days by means of a special adhe- 
sive gland on the head before they begin to feed. Growth is rapid for one to 
three years with juveniles growing to about 6 inches TL by the end of their 



Family and Species Accounts 161 




.V"' < 



Y\ 



Northern Pike (closed circles) and Tiger Muskel- 

lunge (open circles), based on stocking records. 



first summer. When they reach sexual maturity, 3* , ; y p '"' ^ , a 
lengthwise growth slows, but weight increases as the . <'^, M : 

adults become more deep-bodied. Males become sexu- 
ally mature in two or three years, while females mature in three or four 
years. Growth rates vary with the sex of the fish, length of the growing sea- 
son, water temperature, and food availability. Young pike feed on large zoo- 
plankton and immature aquatic insects for three to four weeks before they 
begin to eat small fishes. As top carnivores, adult Northern Pike are vora- 
cious and highly specialized predators that consume a wide variety of prey, 
ranging from fishes to birds and small mammals. 






distribution. Northern Pike are not native to Massachusetts but were 
introduced into the Connecticut River in Vermont and New Hampshire by 
the mid- 1800s. They were first observed in the Connecticut River in Massa- 
chusetts by 1846. These early introductions met with little success, except 
for a small population that became established in the Easthampton Oxbow 
of the Connecticut River. Introductions of Lake Champlain stock in the 
early 1950s established populations in a number of western Massachusetts 
ponds, including Cheshire Reservoir and Onota Lake. Since that time, 
Northern Pike, and more recently Tiger Muskellunge, have been routinely 
stocked statewide as a sport fish and as a management tool to control over- 
abundant or stunted forage fish populations. 

notes . The Tiger Muskellunge is a sterile hybrid between Northern Pike 
and Muskellunge, Esox masquinongy. An overall color pattern of light spots 
on a dark background separates the Northern Pike from the Tiger Muskie, 
which has a vertically striped pattern on the body. 



162 Inland Fishes of Massachusetts 



references. Crossman and Casselman 1987 (bibliography); Raat 1988 
(synopsis); Oatis and Lindenberg 1980 (management in MA); Anon. 1846, 
Storer 1846 (introductions). 



Chain Pickerel Native 

Esox niger Lesueur 1818 plate 37 




identification. Chain Pickerel have fully scaled cheeks and gill covers; 
long, slightly concave snouts; eight or fewer total submandibular pores; 
and a nearly vertical bar beneath the eye. The pattern of narrow, black lines 
against a bright brassy to pale green background forms a chainlike, reticu- 
lated pattern. 

selected counts. D 14-15; A 1 1-13; Scales 117-135; Branchiostegals 
(14)15(17); Submandibular pores 4(5). 

size . Chain Pickerel are intermediate in size between the smaller Redfin 
Pickerel and the larger Northern Pike and Muskellunge. In Massachusetts, 
the average length of three-year-old Chain Pickerel is 13 inches TL, but 
lengths of 15 to 19 inches (1 to 2 pounds) may be attained in more produc- 
tive waters. Older individuals may reach 24 inches and weigh 3 to 4 pounds. 
The Massachusetts state angling record is a 9-pound, 5-ounce (29.5 inches 
TL) fish taken through the ice from Laurel Lake (Berkshires) in 1954. 

natural history. Chain Pickerel occur in a wider range of habitats than 
other esocids and may even be found in brackish waters with salinities up 
to 23 parts per thousand. They typically live in ponds and quiet backwaters 
of medium to large rivers and are usually less common in smaller streams 
inhabited by Redfin Pickerel. The onset of spawning is variable with latitude 
and spring conditions; however, Chain Pickerel move into spawning areas 
after the ice melts and begin to spawn when water temperatures approach 



Family and Species Accounts 163 




«** 







wn localities ^df.-v •Tic- *Ci \ ! ) Jl V 



Chain Pickerel: open circles indicate known localities •;'«(_,- '/ •? •• x 4 %«*l dj% 
where species was not found during our post- 1969 
surveys; not all solid circles were resurveyed. 



1 r*». 



♦\ 






■■•• 



47° to 52°F. In Massachusetts, spawning usually oc- JL 

curs from March to May. Chain Pickerel lay yellowish 
eggs in glutinous strings (up to 9 feet long) in swampy, marshy, or flooded 
areas with abundant submerged aquatic vegetation. Eggs hatch in 6 to 
12 days, depending on the water temperature. After hatching, the fry do not 
feed for six to eight days while they absorb the yolk sac. Chain Pickerel may 
live at least eight years. Food habits and feeding behavior are similar to 
those of other esocids. Juveniles feed on smaller invertebrates and fishes, 
but the adults become highly piscivorous as they grow. Larger Chain Pick- 
erel will eat small mammals, frogs, and snakes. 



■-■:.. /..-■ 



distribution. Chain Pickerel are generally common and widely distrib- 
uted statewide in Massachusetts, occurring in suitable habitats within all 
major drainages. We are not sure if they were naturally distributed on Nan- 
tucket and Martha's Vineyard prior to stocking. 

notes. The Chain Pickerel is an important, native, warmwater game fish 
in Massachusetts due to its widespread occurrence, relatively large size, 
and year-round feeding behavior. Like pike, pickerels are often taken 
through the ice during the winter months. 

references. Crossman and Lewis 1973 (bibliography); Wich and Mull an 
1958 (life history, ecology); Crossman 1978 (taxonomy); Rand and Lauder 
1981 (prey capture) ; Raney 1955 (hybridization); Fiske et al. 1968 (salinity). 



164 Inland Fishes of Massachusetts 



Mudminnow Family 

Umbridae 



The Mudminnow family is a small group of fishes closely related to the 
pickerel and pike (Esocidae). The five species of umbrids have small, non- 
overlapping distributions that pose interesting zoogeographic questions 
about widely separated but closely related species. One species, the Alaska 
Blackfish, Dallia Pectoralis, is found only in western Alaska and another, 
the Olympic Mudminnow, Novumbra bubbsi, is limited to a small area of 
the Olympic Peninsula in Washington. The remaining species are placed in 
Umbra, with the Eastern Mudminnow, Umbra pygmaea, found on the East 
Coast of the United States, the Central Mudminnow, Umbra limi, native to 
the Great Lakes region, and U. krameri found in eastern Europe. With the 
exception of the Alaska Blackfish, mudminnows are small, usually less than 
four inches in length, and have stout bodies. They have nonprotrusible up- 
per jaws and lack spines in their fins. Mudminnows breathe atmospheric 
oxygen using a modified swim bladder, which enables them to survive in 
habitats that become seasonally anoxic. 

references. Cavender 1969, Wilson and Veilleux 1982, Martin 1984 (fos- 
sils, osteology, larvae, and relationships). 



Central Mudminnow introduced 

Umbra limi (Kirtland 1 840) plate 29 




identification. Central Mudminnows are stout-bodied, with rounded 
and opposed dorsal and anal fins that are set far back on the body. They re- 
semble killifishes (Fundulidae), but mudminnows lack a protrusible upper 
jaw and the groove between the snout and the upper jaw. A dark vertical 
bar is usually found at the base of the tail. 



Family and Species Accounts 165 







Central Mudminnow. 






selected counts. D 13-15; A 7-10; 
Scales 34-37. 

(j^k*.* _-) 

size . Central Mudminnows are small fish, usually less 

than 5 inches TL. The largest Massachusetts specimen that we have seen 

measured 98 mm SL. 



//'• 
£v^ 



natural history. The natural history of the Central Mudminnow in 
Massachusetts has not been studied, but, in other areas, they prefer vege- 
tated waters that have little or no current. Small, sluggish streams and quiet 
bays of lakes are common habitats, and in periods of high water, mudmin- 
nows readily swim into areas of submerged terrestrial vegetation. During 
the late spring to early fall, Central Mudminnows are closely associated 
with the vegetation and are most active in the early mornings and late 
evenings. In the cooler periods of the year, individuals generally move out 
of the shallows and into the deeper waters. In some areas, mudminnows 
are active throughout the winter, but this species is reported to burrow into 
the substrate and aestivate (a form of dormancy) during the warmest parts 
of the summer. Spawning occurs in mid- to late spring and is apparently 
associated with high water levels and a rise in water temperature. Central 
Mudminnows may migrate upstream short distances to find suitable 
spawning areas. Females have been found to carry up to 2,000 eggs. Their 
diet is varied, but they frequently take small invertebrates, particularly am- 
phipods and aquatic beetles, as well as small fishes and vegetation. 

distribution and abundance. In Massachusetts, Central Mudmin- 
nows have been found in only a few locations near the Connecticut River 
between Sunderland and Longmeadow. The introduction of this species to 



166 Inland Fishes of Massachusetts 



Massachusetts most likely resulted from the release of laboratory specimens 
from the University of Massachusetts, Amherst, during the mid-1960s. The 
exact date of the introduction is unknown, but the first specimens were 
taken in the wild in 1975. 

notes. Central Mudminnows are able to breathe atmospheric oxygen and 
to survive in waters where other fishes are excluded. It has been estimated 
that one-quarter to one-third of its usual oxygen consumption occurs 
through aerial respiration, which increases during adverse conditions. 
Similar to some other fishes that breathe air, the mudminnow gulps air and 
forces it into its highly vascularized swim bladder, which functions as a 
lung. Air breathing is also enhanced by adaptations of the blood vessels 
that carry blood more directly and more efficiently to the heart. 

references. Chapman 1934 (osteology); Chilton et al. 1984 (winter feed- 
ing); Colgan and Silburt 1984, Martin- Bergmann and Gee 1985, Peckham 
and Dineen 1957, Tonn and Paszkowski 1987 (ecology); Gee 1980 (air 
breathing). 



Family and Species Accounts 167 



Smelt Family 

Osmeridae 



The smelt family consists of seven genera and about 13 species. Found in 
the northern oceans, they are mostly marine or anadromous, but some live 
their full life in freshwater. Smelt are closely related to the salmon and 
trout. Only two species are found in the western North Atlantic, Rainbow 
Smelt and Capelin, Mallotus villosus. Capelin are strictly marine and do not 
range as far south as Massachusetts. Smelt are characterized by an adipose 
fin, a slender body form, and numerous teeth on most of the bones in the 
mouth. Freshly caught members of the smelt group are said to have a dis- 
tinct odor, like fresh cucumbers. Whatever the scent, it is one of the charac- 
teristics that have been used to unite this group of interesting and impor- 
tant fishes. 

references. Hearne, 1984 (development and relationships); Johnson 
and Patterson 1996 (relationships), McAllister, 1963 (review); McDowall 
etal. 1993 (odor). 



Rainbow Smelt Native 

Osmerus mordax (Mitchill 1814) plate 43 




identification. Rainbow Smelt have adipose fins, a slender body, fangs 
on the tongue, relatively large scales (62 to 72 in lateral series), and a long 
lower jaw that reaches to the rear edge of the eye. They lack a well-devel- 
oped pelvic axillary process (see family key Figure 17a, page 56). Smelt are 
silvery; the dorsolateral body is transparent olive-green with an iridescent 
blue to purple sheen. 



168 Inland Fishes of Massachusetts 








W0, 



Rainbow Smelt: open circles indicate stocked popu- v C - %/'\j» % t ,\ cf \ 
lations; coastal records are not indicated. 4/* I fS 



SELECTED COUNTS. D 8-1 1; A 12-16; 
Scales 62-72; GR 26-35. 



M 






size. Most Rainbow Smelt are 7 to 9 inches TL, but occasional specimens 
may reach 13 to 14 inches TL. 

natural history. Anadromous Rainbow Smelt enter the lower edge of 
freshwater to spawn. Unlike salmon, smelt do not pass over dams or stream 
obstructions more than 2 feet high. Smelt spend much of the warmer 
months offshore but apparently never more than three miles from the 
shoreline. During the autumn, adults move back into the estuaries in prepa- 
ration for their spawning migration. In the streams south of Cape Cod, this 
takes place in late February, but in the Gulf of Maine, the runs do not start 
until mid-March. Older, larger fish enter the spawning run first when water 
temperatures reach 39° to 43°F. Spawning is nocturnal, with peaks of 
spawning activity possibly coinciding with bimonthly spring tides. Spawn- 
ing habitat is characterized by gravel and boulder substrate and relatively 
fast-flowing water. Each night, males swim upriver to the spawning sites, 
which are generally in areas where the water velocity is highest. Females 
can produce 7,000 to 45,000 eggs depending on body size. Smelt do not 
build nests; eggs and sperm are broadcast in the water column over a wide 
area by the flowing water. The fertilized eggs attach to the substrate or to 
vegetation and hatch in 11 to 31 days, depending on the water temperature. 
Most of the eggs hatch at night, and the young are swept into the estuary 
where they remain for the first year and grow to a total length of 2 to 5 inches. 
Voracious predators, the smelt feed on crustaceans, insects, worms, and 



Family and Species Accounts 169 



small fishes. Adult smelt are often taken by salmon, trout, Striped Bass, 
Bluefish, birds, and harbor seals. 

distribution and abundance. Historically, Massachusetts Rainbow 
Smelt likely entered almost any unobstructed stream to spawn, but their 
distribution and abundance has been reduced since the turn of the century. 
D.H. Storer remarked that in the mid- 1800s "750,000 dozen smelt" were 
taken annually with dip nets from the Charles River at Watertown. Smelt 
are still found in the lower Charles but, as in other streams, not in the num- 
bers that Storer indicated. Smelt have been introduced into a number of 
inland lakes, ponds, and reservoirs as forage for trout and salmon. This 
practice started many years ago, when Frances Barnard (governor of the 
Bay Colonies, 1760-1769) made a successful introduction of smelt into 
Jamaica Pond, Boston. More recent introductions include the Quabbin, 
Wachusett, and Littleville reservoirs, Mattawa and Wallum lakes, and Big 
Alum, Cliff, Higgins, and Walden ponds. Spawning takes place in some of 
these waters. However, many inland populations appear to be declining, 
apparently due in part to the impacts of acid precipitation. 

notes. Declines in anadromous smelt are primarily due to damming and 
siltation. Dams placed too close to the salt wedge may cause mass egg mor- 
tality due to high salinity and a fungus that results from overcrowding be- 
low the dams. Sport fisheries exist in many areas including the Neponset, 
Fore, Back, and Weir rivers that are tributaries to Boston Harbor. On the 
North Shore, runs are found in the Danvers, Saugus, Annisquam, Parker, 
Rowley, Essex, and Mill rivers. Large runs are also still found in the Jones 
and Weweantic rivers. However, none of these fisheries attracts the esti- 
mated 2,300 smelt fishermen as reported in Boston Harbor by Bigelow and 
Schroeder (1953). 

references. Bigelow and Schroeder 1953, Clayton et al. 1978 (MA ma- 
rine); Murawski et al. 1980 (spawning, Parker River); Murawski and Cole 
1978 (population, Parker River); Reback and DiCarlo 1972 (distribution); 
Storer 1840 (Jamaica Pond introduction). 



170 Inland Fishes of Massachusetts 



Salmon, Char, and Trout Family 

Salmonidae 



The family Salmonidae contains about 70 recognized species and is divided 
into three groups: the salmon, trout, and chars (Salmoninae), the graylings 
(Thymallinae), and the whitefishes (Coregoninae), although the whitefishes 
are sometimes placed in their own family Coregonidae. The Salmoninae, 
the only members of this family found in Massachusetts, have soft-rayed fin 
supports, a dorsal adipose fin, an axillary process at the base of the pelvic 
fins, and a maxilla bone that forms most of the margin of the upper jaw. 
They also have over 100 fine scales in the lateral-line series, well-developed 
teeth on the jaws, and other head bones. 

Since 1875, attempts have been made to introduce or translocate at least 
ten species of salmonids in Massachusetts, with only the exotic Brown Trout, 
Rainbow Trout, and Lake Trout now well established. With the exception of 
the Rainbow Trout, none of the other Pacific salmon, genus Oncorhynchus, 
has met with much success. There are reports of limited reproduction of 
Sockeye "kokanee" Salmon, O. nerka, in Laurel Lake, Lee, and Lake Onota, 
Pittsfield, and there may be a few nonreproducing individuals of Coho 
Salmon, O. kisutch, in the North River from stocking efforts between 1968 
and 1987 that resulted in limited, but documented, reproduction and 
smoking. 

All salmonids reproduce in freshwater and need cold, highly oxygenated 
water, minimal levels of pollution, and silt-free rocky or gravel substrate for 
successful spawning. Reproduction in salmonids is quite similar among 
most species. During spawning, males develop a hooked lower jaw. Ripe fe- 
males select nest sites, usually on gravel-bottomed riffles, and a male de- 
fends the area and stays in close contact with the female. A nest (redd) is 
dug by the female by flapping vigorously with her caudal fin. After nest 
building is complete, the female is joined by the male, and eggs and sperm 
are deposited on the nest and covered with gravel. This process is repeated 
several times over a period of a week or more, and each female may spawn 
with several different males. The eggs remain buried over the winter (in 
fall-spawning species), and hatching occurs from early to late spring. The 
young remain buried in the gravel, absorbing the yolk sac, before emerging. 
Juvenile trout and salmon, called "parr," have a distinctive series of dark 



Family and Species Accounts 171 



blotches along their sides. The blotches, called "parr marks," disappear as 
the fish grows. 

references. Lauder and Liem 1983, Smith and Stearley 1989, Sanford 
1990 (relationships); Maitland et al. 1981 (conservation); Balon 1980 (re- 
view, char); Garman 1895, Cardoza et al. 1993 (introductions); Hearn 1987 
(competition); Halliwell 1989 (local habitat and distribution). 



Key to Adult Massachusetts Salmon, Chars, and Trout 



la. Usually fewer than 12 anal rays 
(rarely 12), length of the fin base shorter 
than longest ray. Go to 2. 

lb. Usually more than 13 anal rays 
(rarely 12 or 13), length of the fin base 
longer than longest ray. Typical Pacific 
salmon (Coho, Chinook), Oncorhynchus. 
See family account. 




2a. Black spots on head and body; pelvic 
and anal fins lacking a white leading edge; 
head and shaft of vomer (midline bone in 
roof of mouth) fully toothed. Go to 3. 

2b. Light spots, usually pink, red, or yel- 
lowish but not black, on head or body; 
leading edge of pelvic and anal fins pure 
white; teeth on head of vomer only. Go 
to 5. 





3a. Caudal fin with radiating rows of 
black spots; red spots never present on 
body; adipose fin with black margin. 
Rainbow Trout, Oncorhynchus mykiss, 
page 174. 

3b. Caudal fin usually unspotted, lacking 
radiating rows of black spots; reddish 
spots sometimes present on body; adi- 
pose fin lacking black margin (not illus- 
trated). Go to 4. 








1 72 Inland Fishes of Massachusetts 



4a. Gill cover usually with several small 
spots; end of jaw reaching last half of eye 
in small fish and beyond the eye in large 
fish; caudal fin broad and usually not 
forked; vomerine teeth well developed. 
Brown Trout, Salmo trutta, page 179, 
Plate 41. 

4b. Gill cover usually with 2 to 3 large 
spots; end of jaw seldom reaching past 
center of eye, except in large males; cau- 
dal fin shallowly forked; vomerine teeth 
not well developed. Atlantic Salmon, 
Salmo salar, page 176, Plate 40. 





5a. Caudal fin weakly forked; body with 
red and white spots; pelvic and anal fins 
with white leading edge followed by a 
contrasting black stripe. Brook Trout, 
Salvelinus fontinalis, page 181, Plate 42. 



5b. Caudal fin strongly forked; sides with 
silver-gray spots; red spots lacking; ante- 
rior edge of pelvic and anal fins not fol- 
lowed by a contrasting black stripe. Lake 
Trout, Salvelinus namaycush, page 183. 




Family and Species Accounts 173 



Rainbow Trout introduced 

Oncorhynchus mykiss (Walbaum 1792) 







identification. Rainbow Trout can be distinguished from the other 
local salmonids by the presence of many small dark or black spots on the 
caudal fin. Adults have a broad pink or red band along their sides and lack 
vermiculations on the back. Juveniles have an olive adipose fin with a black 
margin that is sometimes spotted and a series of 9 to 13 dark parr marks 
along each side. 

selected counts. D 10-13; A 8-12; Scales 100-150; GR 15-16. 

size . Most large Rainbow Trout caught by anglers in Massachusetts 
streams originate from hatchery stocks that average 9 to 12 inches TL when 
released in the spring. Rainbow Trout ranging from 12 to 15 inches TL are 
also stocked in suitable ponds and lakes statewide, and they may grow to 
5 to 6 pounds. The current state record is a 13.3-pound fish taken at John's 
Pond on Cape Cod in 1993. 

natural history. Rainbow Trout are the only spring- spawning salmo- 
nids reproducing in Massachusetts streams. The spawning period extends 
from March to May, coincident with rising water temperature. This species' 
feeding habits, spawning requirements, size, and growth are similar to those 
of native Brook Trout and introduced Brown Trout. Rainbow Trout prefer 
water temperatures between 65° and 68°F. Their distribution in Massachu- 
setts may be limited by their relative intolerance to acidic waters. Repro- 
ducing populations in Massachusetts streams are restricted to cold-water 
streams with a high gradient (more than 75 feet per mile). In these streams, 
they prefer swifter currents, feed more during the day, and are not as secre- 
tive as Brown Trout. Rainbow Trout, particularly hatchery stock, demon- 



174 Inland Fishes of Massachusetts 







"1 Y, Csy'/Y ,Y -} ! \ ) ' < \\\ O) : 



-V >"^, . - 



Rtdnbow Trout: closed circles show reproducing 
populations; open circles indicate localities where 
stocked fishes may survive the summer but not 
reproduce. 



strate greater migratory tendencies than either Brook 
or Brown Trout. Hatchery stocks of Rainbow Trout in 
streams and rivers usually do not hold over, as evidenced by their low re- 
covery rate during summer stream survey work. However, Rainbow Trout 
may hold over in lakes with sufficient cold-water habitat. 




distribution and abundance. Rainbow Trout are native to most of 
the Pacific drainages of western North America and are also found in parts 
of Siberia. They were first introduced into Massachusetts in 1883 and were 
stocked as fingerlings until the 1940s. Today, catchable- sized fish are stocked 
statewide. Reproducing populations of Rainbow Trout are restricted to a 
dozen or so streams in the Connecticut, Deerfield, Westfield, and Housa- 
tonic river drainages. In Massachusetts, coexisting and reproducing popu- 
lations of Brook, Brown, and Rainbow Trout occur only in a single tributary 
to the Housatonic River in Lanesborough. 

notes. Rainbow Trout were formerly treated under the species, Salmo 
gairdneri, which erroneously indicated a relationship with the Atlantic 
salmon group. Recently, the North American Rainbow Trout has also been 
shown to be the same species as the Rainbow Trout of the Kamchatka Pen- 
insula in Siberia. Thus, it was necessary to change the specific name to my- 
kiss. Later it was documented that the Rainbow Trout are more closely re- 
lated to the Pacific genus Oncorhynchus than to the Atlantic genus Salmo so 
that now the American Fisheries Society recommends the use of Oncorhyn- 
chus mykiss as the correct name. 



Family and Species Accounts 1 75 



references. Smith and Stearley 1989 (taxonomy); Halliwell 1989 (distri- 
bution, habitat, MA); Simmons 1997 (summer trout). 



Atlantic Salmon Native stocks extirpated 

Salmo salar Linnaeus 1758 plate 40 



identification. Atlantic Salmon and the related Brown Trout have 
teeth on the head and shaft of the vomer. Adult Atlantic Salmon have 
poorly developed deciduous teeth on the shaft of the vomer (well devel- 
oped in Brown Trout) and small X-shaped spots on the body. Atlantic 
Salmon also have a smaller mouth, a more deeply-forked tail, and longer 
pectoral fins. They can be distinguished from the members of the Pacific 
salmon genus Oncorhynchus by the lack of black spots on the caudal fin. 
Juvenile Atlantic Salmon (parr) have 8 to 1 1 narrow parr marks with a single 
red spot between each pair of parr marks. 

selected counts. D 10-13; A8-11; Scales 110-120; GR 15-19. 

size. The average size of anadromous and landlocked Atlantic Salmon dif- 
fers, with landlocked populations usually reaching only 20 inches TL. Anad- 
romous forms commonly reach 30 inches TL. In anadromous forms, ocean 
growth is rapid and maximum size is larger; males returning after one year 
at sea weigh 3 to 6 pounds, fish returning after two years range from 6 to 
12 pounds. Repeat spawners may weigh up to 40 pounds. In contrast, land- 
locked salmon average 2 to 4 pounds. The Massachusetts state record land- 
locked Atlantic Salmon, angled from Wachusett Reservoir in 1985, weighed 
10 pounds, 2 ounces. 

natural history. Anadromous Atlantic Salmon spawn in freshwater 
streams and then return to the sea. Historic accounts show that they mi- 



176 Inland Fishes of Massachusetts 



. -7 - 




Atlantic Salmon: closed circles show areas stocked 
with juveniles or returning adults from anadromous 
restoration program; open circles indicate landlocked 
reproducing populations. 



U' 



/;■■ 



grated far into New Hampshire and Vermont in the ;J 9 * J ~> v 

Connecticut River and well into New Hampshire along the 
Merrimack River. Young salmon remain in freshwater for two to three 
years, descending to the sea as "smolts" when they reach 5 to 9 inches TL. 
At sea, they live for one or two more years before returning to their natal 
streams to spawn. Although spawning occurs from October through No- 
vember, most adults return to freshwater well before then, typically in 
May and June (see family account for general spawning habits and require- 
ments). Unlike most Pacific salmon, which die after spawning, many post- 
spawning Atlantic Salmon survive and return to the sea. Food habits vary 
with life stages. At sea, salmon eat a variety of marine organisms, includ- 
ing crustaceans and smaller fishes. Despite their willingness to accept an 
angler's fly, adult Atlantic Salmon do not feed in freshwater prior to spawn- 
ing. Young Atlantic Salmon feed primarily on aquatic and terrestrial insects 
while they are in freshwater. Landlocked Atlantic Salmon in large Massa- 
chusetts reservoirs feed principally on introduced Rainbow Smelt, young 
White Perch, and midges and ants (J. Bergin 1998, pers. comm.). 






distribution and abundance. In Massachusetts, native anadromous 
Atlantic Salmon were historically known from the Connecticut and Merri- 
mack rivers. Populations may also have been present in other suitable rivers 
before they were overfished or dammed, but there is little information that 
is not anecdotal. Early accounts mention their great abundance, particu- 
larly in the Merrimack, where 60 to 100 were taken daily near the river mouth 
in 1790. The Connecticut River had a large run until dams constructed at 



Family and Species Accounts 177 



Hadley (1794) and Turners Falls (1798) eliminated the upstream salmon 
run in as little as 12 years. Additional dam construction at Lawrence on the 
Merrimack River in 1847 destroyed one of the finest Atlantic Salmon runs 
in New England. Stolte (1981) estimated that the historic population in the 
Merrimack Drainage possibly ranged from 8,940 to 26,820 adults. 

For many years, ongoing attempts to restore Atlantic Salmon to Massa- 
chusetts rivers have had limited success due to a combination of many fac- 
tors, including poorly designed fishways, inferior genetic stock, turbine 
mortality, and poor survival rates at sea. Following the release of millions 
of fry and smolts over a 20-year period, the first redds and eggs of Atlantic 
Salmon were found in the main stem of the Westfield River by biologists 
J. O'Leary and D. Pugh in November 1994. Since then, a few additional 
redds have been found in upper tributaries to the Westfield River. Land- 
locked Atlantic Salmon have been introduced into the Quabbin, Wachusett, 
and Littleville reservoirs. These landlocked forms, originating from lakes in 
Maine and New Hampshire, are genetically similar to anadromous Atlantic 
Salmon but differ primarily in their nonmigratory habits and their ability to 
live in deep lake environments. 

notes. Landlocked salmon were first introduced into Quabbin Reservoir 
in May 1965, with 14,400 spring yearlings from Maine. Most of the adult 
anadromous Atlantic Salmon that are now captured at fish lifts at the Hol- 
yoke Dam on the Connecticut River and the Lawrence Dam on the Merri- 
mack River from May to June were stocked as yearlings (Figure 3). Most of 
these adult salmon are captured and artificially spawned under hatchery 
conditions, and their young are released into selected tributaries during the 
following spring. 

references. Bridges andHambly 1971, Bergin 1996 (Quabbin Res.); So- 
chasky 1981, Lacroix et al. 1985 (impacts of acidification); Rosseland and 
Skogheim 1984, White et al. 1984 (acid mitigation); Gibson 1973, Symons 
1974, Kennedy and Strange 1980, Hearn 1987 (competition); Saunders 
1981 (management); Behnke 1972, 1986, 1988 (taxonomy); Kendall 1935; 
Dymond 1963 (biology, historic records); Stolte, 1981 (Merrimack popula- 
tion); Rideout 1989, Rideout and Stolte 1988, Foster 1991, O'Leary 1995 
(restoration). 



1 78 Inland Fishes of Massachusetts 



Brown Trout introduced 

Salmo trutta Linnaeus 1 758 plate 41 



identification. Bro wn Trout and the related Atlantic Salmon have 
well-developed teeth on both the head and on the shaft of the vomer, the 
bone along the midline of the roof of the mouth. Brown Trout have a large 
mouth that reaches well beyond the eye in large fish (to about mideye in 
6-inch fish) and usually have an almost square tail. The sides, back, and 
dorsal fin have dark brown-black spots, often framed in a halo of blue. Red- 
orange spots are often found on the adipose fin and sides, usually below 
the lateral line. They lack white edges on the pelvic and anal fins and black 
spots on the caudal fin, and generally lack the Atlantic Salmon's X- shaped 
markings on the sides. 

selected counts. D 10-14; A 8-12; Scales 120-130; GR 14-18. 

size . Massachusetts Brown Trout typically range from 8 to 10 inches TL, 
although older fish often grow to 15 inches TL. Brown Trout inhabiting 
cold-water ponds and lakes, as well as coastal "sea-run" populations, grow 
larger and weigh up to 10 pounds. The current state record for Brown Trout, 
a 19-pound, 10-ounce fish, was taken in 1966 from Wachusett Reservoir. 

natural history. Brown Trout spawn from late autumn to early winter 
(October to December) in tributary streams and small rivers. They prefer 
spawning substrate with stones ranging from 0.25 to 3 inches in diameter. 
Larger females tend to select areas with larger stones. Stream populations 
of Massachusetts Brown Trout generally live longer and grow faster than 
Brook Trout. Most stream-dwelling Brown Trout mature at the end of their 
second year of life and generally live for at least three growing seasons. They 
rarely live more than five years in the wild. Juvenile Brown Trout are pri- 
marily insectivorous, but as they approach 8 to 12 inches TL, their diet shifts 



Family and Species Accounts 179 




MX 

Brown Trout: closed circles show reproducing popu- : ( (j / -L/ 
lations; open circles indicate localities where stocked 
fishes may survive the summer but not reproduce. 



to more fish. Like Brook Trout, Brown Trout re- 
quire highly oxygenated cold water and often compete 
with Brook Trout for food and habitat. However, Brown 
Trout can tolerate higher water temperatures than Brook Trout. In cold- 
water habitats, the largest Brown Trout are found in lower stream reaches 
where cover and large, deep pools are found. 




distribution and abundance. Brown Trout, natives of Europe and 
western Asia, were first introduced into Massachusetts in 1887. Original 
stocks included both the German Brown strain from central Europe and the 
Loch Leven strain from Scotland. Today, reproducing Brown Trout occur in 
most cold-water streams of the Berkshire Valley and Central Uplands of 
Massachusetts, but they are only sporadically found farther east. Sea- run 
populations are reported from a few of the larger streams in the Falmouth - 
Mashpee area of Cape Cod. Aside from stream gradient and temperature, 
a major factor limiting the distribution of Brown Trout in Massachusetts 
is acidification. Brown Trout will not thrive in areas that do not have the 
capacity to naturally buffer acids. 

notes . The first Massachusetts stocking policies limited the introduction 
of Brown Trout to waters where they would not compete with the native 
Brook Trout. However, the issue of competition between the two species 
was forgotten, and Brown Trout were introduced into areas with estab- 
lished Brook Trout populations. The original stocking policies for Brown 
Trout were valid, because Brown Trout can negatively influence Brook 
Trout populations. Consequently, we believe new introductions of Brown 



180 Inland Fishes of Massachusetts 



Trout should not be made where native populations of Brook Trout exist in 
the absence of Brown Trout. 

references. Belding 1920 (introduction, MA); MacCrimmon and Mar- 
shall 1968 (worldwide distribution); Fausch and White 1981, Hearn 1987 
(competition); Bachman 1984 (foraging behavior); Horton 1961, McFadden 
and Cooper 1962 (population ecology); Bergin 1984 (sea-run stock manage- 
ment); Halliwell 1989, Simmons 1997 (distribution and macrohabitat, MA). 



Brook Trout or Brook Char Native 

Salvelinus fontinalis (Mitchill 1814) plate 42 



identification. The chars (Brook and Lake Trout) lack teeth on the 
shaft of the vomer and black spots on the body. Brook Trout have heavy 
dorsal vermiculations (worm-like markings) and a dark stripe behind the 
white leading edges of the pelvic and anal fins. In addition, they have a 
squarish or shallowly forked tail and red spots, often with blue halos, along 
the sides. Colors intensify during spawning and the lower flanks and belly 
of males become deep magenta. 

selected counts. D 10-14; A 8-13; Scales 110-130; GR 14-22. 

size . Adult Brook Trout in Massachusetts streams reach 6 to 8 inches TL, 
but total lengths of 10 to 12 inches are possible in unexploited populations. 
One- or two-year-old fish ranging from 3 to 6 inches TL are most numerous 
in stream populations. Brook Trout living in cold-water ponds and lakes and 
the coastal Salter populations (a sea-run form) grow considerably larger 
and faster. The current state record for a Brook Trout is 6 pounds, 4 ounces; 
it was caught in 1968 from Otis Reservoir in western Massachusetts. 



Family and Species Accounts 181 




Brook Trout: closed circles show current repro- 
ducing populations; open circles indicate localities 
where stocked fishes may survive the summer but not 
reproduce. 



SU-' 



dry 



\ 






natural history. In Massachusetts, Brook Trout 
inhabit flowing, highly oxygenated, cold-water streams. 
They tolerate a variety of habitats, from high-gradient mountain streams to 
low-gradient meadow brooks generally kept cool by groundwater or springs. 
The sea-run form, or Salter, has a life cycle similar to that of the Atlantic 
Salmon, with adults spending part of their lives in salt water. Brook Trout 
have more rigid temperature requirements than do Brown Trout, Rainbow 
Trout, or Atlantic Salmon. They generally do not tolerate water tempera- 
tures exceeding 68°F for extended periods of time. Studies in Massachusetts 
indicate that the optimum range for maximum activity and feeding is be- 
tween 55° and 65°F. Brook Trout spawn in both lakes and streams, although 
lake-spawning populations are rare in Massachusetts. Spawning lasts from 
late September through November, but elevation and water temperature 
influence the exact timing. Stream populations spawn over gravel riffles 
composed of coarse sand or stones up to 4 inches in diameter. Individuals 
mature at an early age, and some males capable of spawning during their 
first year are between 3.5 and 4 inches, although most males mature during 
their second year. The life span of Brook Trout in Massachusetts streams 
seldom exceeds three growing seasons. Stream Brook Trout are insectivo- 
rous throughout their lives. Pond-dwelling and Salter Brook Trout popula- 
tions tend to consume fish in addition to invertebrates. 



>Zs< 



distribution and abundance. Reproducing Brook Trout are found 
all across Massachusetts. Due to widespread introductions, native popula- 
tions are difficult to distinguish from stocked populations. Brook Trout are 



182 Inland Fishes of Massachusetts 



most common in western and central Massachusetts and occur only spo- 
radically in the east. The eastern populations have declined, and today only 
a few geographically isolated populations remain. Likewise, Salter Brook 
Trout that were historically found in one or two tributaries to Massachu- 
setts Bay have been extirpated. Salters, though reduced in numbers, are still 
known from a few tributaries to Nantucket Sound, as well as Buzzards and 
Narragansett bays, but these populations have not been studied critically. 

notes . The remnant Brook Trout populations in eastern Massachusetts 
are indicators of the location of relatively undisturbed environments. Their 
continued presence serves as a barometer for measuring the condition of 
the environment for the trout and other organisms that require cold clean 
water. 

references. Bridges and Mullen 1972 (life history, MA); Bigelow 1963 
(review); Power 1980 (life history); Behnke 1980 (taxonomy); Estes 1987 
(bibliography); Mullan 1958, Ryther 1997 (sea-run, MA); Halliwell 1989 (MA 
distribution, habitat); Wydoski and Cooper 1966 (growth); McFadden 1961 
(population); Daye and Garside 1975, 1976 (acid conditions); Hearn 1987 
(competition). 



Lake Trout or Lake Char introduced 

Salvelinus namaycush (Walbaum 1792) 



identification. The chars (Lake and Brook Trout) lack black spots 
on the body and teeth on the shaft of the vomer. Lake Trout have a more 
streamlined body and a distinctly forked tail and lack the red spots found 
on the sides of Brook Trout. In addition, Lake Trout have more than 92 py- 
loric caeca while Brook Trout have fewer than 56. Body color ranges from 



Family and Species Accounts 183 



/'«' 







Lake Trout. 






silver-gray to almost black dorsally with a silvery K \\\t \ U$ 

white belly and throat. Whitish spots mark the 

body and the dorsal and caudal fins. The pelvic and ,... _/3%j.a^.S 

anal fins sometimes have white leading edges and an 

orange wash, but the markings are not as conspicuous as in Brook Trout. 



N\ 



^< 



selected counts. D 8-11; A 8-10; Scales 116-138; GR 16-26. 

size . The Lake Trout is the largest North American char. Adult Massachu- 
setts Lake Trout average 18 to 27 inches TL and 2 to 8 pounds, respectively. 
The current Massachusetts record, weighing 22 pounds, 10 ounces and 
measuring 38 inches TL, was taken at Quabbin Reservoir in 1988. 

natural history. Lake Trout require large, deep, cold-water lakes; in 
Massachusetts, they are confined to built reservoirs. During the summer, 
adult Lake Trout inhabit depths of 60 to 130 feet and prefer a constant 
water temperature of 55°F. In late fall, winter, and spring, Lake Trout move 
into shallow waters. In Massachusetts, Lake Trout spawn during October 
and November at depths of 10 to 75 feet, over rubble, riprap, or rock sub- 
strates. Two strains, a shallow- water, shoal-spawning form and a deep- 
water- spawning form, are suspected to exist in Massachusetts. Male Lake 
Trout usually do not spawn until they are 14 to 17 inches TL and four to six 
years old. Females first spawn at 18 to 21 inches TL and six to eight years of 
age (J. Bergin 1998, pers. comm.). Individuals spawn over a two -week period 
from sunset to midnight. Males usually precede females to the spawning 
grounds, where they sweep away debris by fanning and rubbing the sub- 
strate. Following the arrival of females, spawning starts and the fertilized 



184 Inland Fishes of Massachusetts 



eggs sink between rocks where they are left unattended. The eggs hatch from 
late February to April, and the young gradually move into deeper waters. 
Lake Trout are long-lived and slow-growing fishes. In November 1970, 
a 27-inch male Lake Trout was tagged and released at Quabbin Reservoir; 
when it was recaptured in July 1985, it measured only 35 inches TL. For the 
first two years of life, Lake Trout feed primarily on small insects and zoo- 
plankton. By 12 to 15 inches TL, they are primarily piscivorous, feeding on 
almost any suitably sized fish found in their deep, cold-water habitats. In 
Massachusetts, they feed primarily on introduced Rainbow Smelt and, to a 
lesser extent, on White Perch, Yellow Perch, and isopods. 

distribution and abundance . While there were some unsuccessful 
Massachusetts introductions of Lake Trout in the 1870s, the first recent in- 
troduction occurred in 1952 with stocks of fingerlings from the Lake On- 
tario region of New York. The later stocks (1965-1970) come from a Finger 
Lakes deepwater strain (Seneca-Cayuga Lakes). Lake Trout are found only 
in the Quabbin and Wachusett reservoirs. Both areas are characterized by 
deep, cold, well- oxygenated water. 

references. Martin and Oliver 1980 (life history); Behnke 1980, 1984 
(taxonomy); Johnson et al. 1987 (acidification-bioassay); Bridges and Ham- 
bly 1971, Keller 1987 (Quabbin management); Marshall and Keleher 1970 
(bibiliography 1929-1969); Daly et al. 1962 (life history); Martin and Bald- 
win 1960 (hybridization). 



Family and Species Accounts 185 



Trout-perch Family 

Percopsidae 



The Trout-perch family is characterized by an unusual combination of both 
primitive and advanced features. An adipose fin is present, as in trout and 
catfishes, and there are true fin spines, an advanced feature found, for ex- 
ample, in sunfishes and perch. This family has a relatively large and slightly 
subterminal mouth, a nonprotrusible upper jaw, small teeth, and ctenoid 
scales. Trout-perch are thought to be closely related to the Pirate Perch 
(Aphredoderidae) and the cavefishes (Amblyopsidae). The Percopsidae is 
a small group comprising a single genus and two species that are endemic 
to North America. One species is widely distributed in eastern and middle 
North America, while the other species is restricted to a portion of the Co- 
lumbia River in the extreme northwestern United States. Trout-perch are 
generally small, usually less than 5 inches in length, and nocturnal. Where 
abundant, Trout-perch are an important forage fish for many of the larger 
game fish, including trout, Yellow Perch, and Northern Pike. 

references. Lauder and Liem 1983 (relationships); Scott and Crossman 
1973 (general). 



Trout-perch Native, State Extirpated 

Percopsis omiscomaycus (Walbaum 1 792) plate 30 




identification. Trout-perch are distinguished from all other Massa- 
chusetts fishes because they have the combination of an adipose fin, true 
spines, a slightly subterminal mouth, ctenoid scales, and lack barbels. They 



1 86 Inland Fishes of Massachusetts 



A tf \ • 



'i . y y 



:b;: 



/v- 



vi V'/ 



v.Vr' SJ/ 



^ 



y?s 



A. 



"'" ' "'■ "]\r " ■■ j s^V; \\'-< -«/" ,v ~ S ^'V 



Trout-perch. 






are silvery-yellow to cream with a series of small, 
dark spots along the upper portions of the body. 



K 



v 



,PSr&J-< 



selected counts. D 11,10-1 1; A 1,6-7; Scales 42-60. 



-... p\ 



size. Trout-perch are small fishes; most individuals are less than 
5 inches TL. 

natural history. Trout-perch were last found in Massachusetts in the 
early 1940s. Information about their natural history in this state is unknown. 
In other parts of their range, Trout-perch are still common and have been 
extensively studied. They typically inhabit lakes and quiet, backwater pools 
in streams. Trout-perch stay in deep water or under cover during the day 
and become active at night. Their diet consists of a variety of small inverte- 
brates, including mayflies, amphipods, midges, and, occasionally, small 
fishes. Spawning occurs from mid- to late spring in small, rocky tributary 
streams or in shallow, inshore areas of lakes. Females have been found to 
carry up to 750 eggs. 

distribution and abundance. In Massachusetts, Trout-perch were 
known from only three localities in the extreme western portion of the state. 
The earliest record is based on a single specimen collected at Williamstown, 
Hoosic Drainage, prior to 1859. Six specimens were collected near the 
mouth of the Green River, Housatonic Drainage, between 1940 and 1942 
(McCabe 1942), but these specimens have been lost. A second collection of 
12 specimens from the Green River, a half-mile above the Housatonic River, 
was also made by B. McCabe, and they are now preserved at the Museum 



Family and Species Accounts 187 



of Comparative Zoology. The Trout-perch may never have been common in 
this state. Surveys between 1978 and 1990 have failed to locate this species, 
and it is now presumed to have been extirpated from Massachusetts. 

notes. As Trout-perch are nocturnal and rather secretive, it is possible 
that populations still survive in Massachusetts. Surveys in the upper Hoosic 
Drainage in New York, downstream of Massachusetts, have found the Trout- 
perch to be common in the mid-1980s. But in the Housatonic Drainage in 
Connecticut, Trout-perch have not been collected since 1879 (Whitworth 
et al. 1968). Extensive pollution in the Hoosic and Housatonic drainages 
may have caused the decline of this species. Extirpation of the Trout-perch 
in Virginia is thought to be due to introduced predators coupled with other 
factors. 

references. Crowder et al. 1981 (diet); Kinney 1950 (life history); Mc- 
Cabe 1942 (first MA record); Magnuson and Smith 1963 (life history); Smith 
1985 (NY records); Jenkins and Burkhead 1993 (VA). 



1 88 Inland Fishes of Massachusetts 



Cod Family 

Gadidae 



The cod family consists of about 30 species that are found primarily in the 
marine waters of the Northern Hemisphere. Cods belong to the order Gadi- 
formes, a group that also contains such fishes as Pollack, hakes, whitings, 
haddock, cusk, and rocklings. The codfish order has been historically char- 
acterized in part by a specialized arrangement of the caudal vertebrae and 
caudal fin rays. However, the group's interrelationships are confusing due 
to many reduced or lost characters that are difficult to interpret. The Bur- 
bot, Lota lota, is sometimes placed in the family Lotidae or the subfamily 
Lotinae. Cods have one, two, or three dorsal fins plus one or two anal fins, 
and most species have a single, median chin barbel. Cods are among the 
world's most commercially important fish families, with about 12 million 
metric tons being taken worldwide each year. 

references. Bigelow and Schroeder 1953 (marine species); Markle 1989, 
Patterson and Rosen 1989 (relationships); Fahay 1983, Dunn and Matarese 
1984 (larvae, development and relationships); Jensen 1972 (cod fishery); 
Cohen 1989 (general). 

Key to the Massachusetts Freshwater Cods 



la. Two dorsal fins; length of the base of 
the second 6 or more times the length of 
the first; one anal fin. Burbot, Lota lota, 
page 190, Plate 39. 



lb. Three dorsal fins; bases of each of 
near equal lengths; 2 anal fins. Atlantic 
Tomcod, Microgadus tomcod, page 192, 
Plate 38. 




Family and Species Accounts 1 89 



Burbot Native, Special Concern 

Lota lota (Linnaeus 1758) plate 39 




identification. Burbot have a median chin-barbel, long second dorsal 
and anal fin, and an elongate body. They are the only member of the cod 
family restricted to freshwater. The upper body is light brown to gray and 
mottled with darker shades of brown and gray. The dorsal and caudal fins 
of young Burbot are usually edged with dark pigment, and a dark spot with 
a light ocellus is on the upper caudal fin. 

selected counts. D 10-12, 66-67; A 65-71. 

size . Burbot may grow as large as 3 feet TL in some parts of their range. 
The largest specimens are usually found in large, deep lakes or rivers. Indi- 
viduals living in smaller rivers and streams grow only to lengths of slightly 
over 12 inches TL. 

natural history. Little is known about Burbot in Massachusetts. In 
other areas, they are a resident of cold deepwater lakes and rivers. Burbot 
enter tributaries in late winter and spring and spawn nocturnally, under the 
ice, between November and March. They spawn in tight groups that re- 
semble a ball of whirling fishes. Post-larvae, recently collected near the 
mouth of Longmeadow Brook, Longmeadow, were probably spawned in 
February. This species is largely nocturnal. During the daytime, juveniles 
can be found under large, flat rocks in a foot or so of water. Young Burbot 
feed on aquatic insects and crustaceans, but large adults feed almost exclu- 
sively on fishes. Their choice of prey depends on availability and some- 
times includes Alewives, salmon, smelt, sculpins, and sticklebacks. 

distribution and abundance. Burbot are distributed throughout 
the Northern Hemisphere. The southernmost population of Burbot in east- 



190 Inland Fishes of Massachusetts 




Burbot: open circles indicate known 
species was not found during our post 
not all solid circles were resurveyed 



u ,;;% , 

localities where ] ( '( / i \, • 4 % g I \J\ >-- 

■r ; i, n r ,\IX< - 
;t- 1969 surveys; ^L/j \j (\ - * 

I 






ern North America is known from the Twin Lakes 
region of Connecticut in the Housatonic River Drain- 
age. In Massachusetts, Burbot are rare; to date, they are known from only 
a few records. Lesueur (1818) described and illustrated a juvenile specimen 
from Northampton (presumably from the Connecticut River). Storer (1839) 
stated that he saw a 6-inch fish from the Ashuelot River, New Hampshire 
(Connecticut Basin), but later (1867) he mentions another (or the same?) 
6-inch specimen from the Connecticut River (in Massachusetts?). In July of 
1970, J. Bergin and a Massachusetts Division of Fisheries and Wildlife sur- 
vey crew found single specimens (7.5 and 8.5 inches TL) in Hubbard and 
Schenob brooks, both in the Housatonic Drainage, near the Connecticut 
state line. The most recent record is based on two post-larvae found on 
May 16, 1987, by W. Kenney, a Springfield aquarist, in Longmeadow Brook 
near the Connecticut River. One of the specimens (now in the MCZ collec- 
tion) was raised until mid-June, when it measured about 1 inch TL (30 mm 
SL) and was definitely identified as a Burbot. The most recent records are a 
1998 MDFWfind of a 12-inch fish at the mouth of the Fort River, Hadley, 
and another 12-inch specimen brought to Douglas Smith at the University 
of Massachusetts by an angler who caught it in a pool below the Turners 
Falls dam on the Connecticut River in December, 2000. The angler noted 
that other specimens have been caught. This most recent record is not plot- 
ted on the distribution map. 



\\ 



notes. Due to the species' rarity, the Burbot is listed as a Species of Special 
Concern in Massachusetts, but its actual status is an enigma. Stream sur- 
veys attempting to locate Burbot in the Housatonic (1979-1989) and in the 



Family and Species Accounts 191 



Connecticut tributaries (1979-1988) failed to find the species. Burbot were 
never collected during hundreds of gillnet sets or seine hauls made over a 
20-year period (1969-1989) by the Massachusetts Cooperative Fisheries Re- 
search Unit in the Connecticut River main stem. In addition, local anglers 
did not report this species until one was brought to our attention in De- 
cember, 2000. Since some Burbot found at the southern edge of their range 
may be vagrants from established populations, it is possible that the Con- 
necticut River fishes are from upriver in New Hampshire. If this is correct, 
the eggs, larvae, juveniles, or prespawning adults would have been washed 
at least 100 miles downstream and over at least three major dams. In con- 
trast, it is also possible that there still exists a small population of Burbot in 
one of the deep pools along the Connecticut River or within its major tribu- 
taries. Whitworth (1996), with limited data, states that the species is intro- 
duced in the Connecticut and Housatonic drainages in Connecticut. 

references. Bergin 1970 (Housatonic records); Lesueur 1817 (first MA 
record); Scott and Crossman 1973, Lee and Gilbert 1980 (general); Storer 
1839 (historical record); Whitworth (1996). 



Atlantic Tomcod 

Microgadus tomcod (Walbaum 1792) 



Native 
plate 38 




identification. Atlantic Tomcod have a chin barbel, three dorsal fins, 
and two anal fins. Its small size, rounded caudal fin, and the filamentous 
extensions of the second pelvic fin ray distinguish it from the marine Atlan- 
tic Cod, Gadus morhua, haddock, Melanogrammus aegleflnus, and Pollack, 
Pollachius virens. 

selected counts. D 1 1-15,15-19,16-21; A 12-21,16-20; GR 16-21. 



192 Inland Fishes of Massachusetts 



..; : 




: r w 



), K -■'*/' 



Atlantic Tomcod: coastal records are not indicated. f (/ j -i, 'J* ^ i ) i^ I J^- 

ly,(\ r 

s i ze. This small species of cod usually reaches * / i/ v 

only 9 to 12 inches TL. There are historic reports of : ,-' x 
15 -inch Atlantic Tomcod, weighing up to 1 .25 pounds. ,J$%.<>m 

natural history. Atlantic Tomcod are most abundant in brackish es- 
tuaries and shallow harbors. They move inshore and upstream from Octo- 
ber to May, and then downstream to slightly deeper, coastal marine waters 
in the late spring. Spawning occurs in shallow water during the coldest 
months of the year, when water temperatures range from 34° to 41°F. Ice 
often provides cover and may contribute to the suitably low salinities re- 
quired during the spawning and incubation period. In the Weweantic River 
estuary in southeast Massachusetts, Atlantic Tomcod spawn between No- 
vember and late January, with peak activity in late December. Juveniles 
reach 3 to 3.5 inches TL by July. Growth rate slows during late summer, but 
as temperatures decline, Atlantic Tomcod resume growth and reach 5 to 
7 inches TL at the end of their first year. Most Atlantic Tomcod mature in 
their second year when they are approximately 10 inches TL. Those inhab- 
iting southern New England estuaries and the Hudson River usually spawn 
only once and seldom live more than two years. Atlantic Tomcod are op- 
portunistic feeders and prey on a variety of copepods, amphipods, and 
decapods, as well as small fishes. 

distribution and abundance. In Massachusetts, Atlantic Tomcod 
can be expected to occur in most coastal areas. They move to the head of 
the tide and above in unobstructed coastal streams during the winter spawn- 
ing season. Storer (1839) noted that 2,000 bushels were taken annually from 
the Charles River at Watertown. Although still common, Atlantic Tomcod 



■ 



...■l.y 



Family and Species Accounts 193 



have declined in some areas. The declines are probably due to changes in 
the water quality of coastal streams, inner harbors, and bays. 

notes. Atlantic Tomcod, sometimes called "frostfishes," are delicious and 
easy to catch. They are often caught by anglers fishing for smelt and will 
take almost any kind of bait. Atlantic Tomcod remains have been found at 
Native American archaeological sites as far inland as Marlborough. The 
species also supported a small, local commercial fishery until about 35 years 
ago. Although Tomcod can tolerate drastic changes in salinity and tempera- 
ture, they seem particularly susceptible to acidification, altered stream flow, 
chemical pollution, and low oxygen levels. Scott and Crossman (1973) state 
that while "...some other fish species may be able to move out of the region 
[of pollution], to return when pollution ceases, the tomcod is unable to re- 
produce and soon disappears entirely from the affected region." 

references. Bigelow and Schroeder 1953 (general); Clayton etal. 1978 
(review); Howe 1971 (life history, Weweantic estuary, MA); McLaren et al. 
1988 (life history, Hudson River); Scott and Crossman 1973 (review). 



194 Inland Fishes of Massachusetts 



Needlefish Family 

Belonidae 



The needlefishes, family Belonidae, are typically found in temperate and 
tropical marine waters worldwide. However, one-third of the belonid spe- 
cies, such as those from the Amazon River, live their complete lives in 
freshwater. The needlefishes belong to a relatively small group of fishes, 
called the Beloniformes, which contain such groups as the flying fishes 
(Exocoetidae), the halfbeaks (Hemiramphidae), and the sauries (Scombere- 
socidae). The needlefish family contains 32 species in 10 genera. Three spe- 
cies are found in Massachusetts marine waters, but only one, the Atlantic 
Needlefish, enters coastal freshwaters. Belonids are elongate fishes with 
opposing dorsal and anal fins set far back on the body. Most adult needle- 
fishes have elongate upper and lower jaws studded with small, sharp teeth. 
The development of the jaws is interesting in a evolutionary sense because 
most needlefishes pass through a stage in which the upper jaw is much 
shorter than the lower jaw. This elongate lower jaw can be found in some 
juvenile flyingfishes and persists in the adults of the halfbeaks. 

references. Collette et al. 1984 (development, relationships); Collette 
and Berry 1965 (nomenclature, systematics); Collette and Parin 1970 (re- 
view, Eastern Atlantic); Cressey and Collette 1970 (copepod parasites). 



Atlantic Needlefish Native 

Strongylura marina (Walbaum 1792) 




identification. Needlefishes have elongate bodies, posterior and op- 
posing dorsal and anal fins, and long forcep-like jaws studded with many 
teeth. The Atlantic Needlefish is the only local species that readily enters 
freshwaters, but two other species, the Flat Needlefish, Ablennes hians, and 



Family and Species Accounts 195 



the Agujon, Tylosurus acus, occur in marine waters. Atlantic Needlefish 
have rounded or slightly indented tail fins while the tails of the other two 
species are deeply forked. In addition, Atlantic Needlefish lack a gonad on 
their right side. 

selected counts. D 14-17; A 16-17; Predorsal scales 213-304. 

size. Adults commonly range to 20 inches TL; however, specimens as large 
as 2 feet TL have been noted (640 mm SL). 

natural history. Atlantic Needlefish are absent from New England 
waters until early spring, when migrants arrive from the south. The year- 
round surveys of the Massachusetts Division of Marine Fisheries found 
them in estuaries between July and October. Adults enter coastal rivers and 
streams to spawn over submerged vegetation in shallow brackish to fresh 
waters. After the eggs are deposited, they attach to the vegetation with nu- 
merous filaments. Young hatch at 0.75 inches TL with neither of the jaws 
elongated. As the fish grows, the lower jaw grows faster than the upper, and 
at 1.5 inches TL, the upper jaw is only one-half the length of the lower. The 
upper jaw remains shorter than the lower until the young reach approxi- 
mately 9 inches TL. Young Atlantic Needlefish feed on small crustaceans, 
particularly shrimp, and switch to small fish such as silversides and killi- 
fishes when they are larger than 3 inches TL. 

distribution and abundance . Atlantic Needlefish have been found 
100 miles up the Hudson River and in the Connecticut River almost to the 
Massachusetts state line. In Massachusetts, Atlantic Needlefish are com- 
mon in the coastal rivers and bays south of Cape Cod. They have been 
recorded from Pleasant and Waquoit bays and in the Bass, Slocums, Ware- 
ham, and Westport rivers. Local residents have told us of large "garfishes" 
in Weir Mill Creek above Follins Pond at the head of the Bass River on Cape 
Cod during spring. Large numbers have been taken in Pleasant Bay on 
Cape Cod, but this area lacks major freshwater tributaries. Based on a 
single larva (0.6 inches TL) collected near the Pilgrim Nuclear Power Plant 
in Plymouth on July 28, 1977, the species has apparently bred north of Cape 
Cod in Massachusetts Bay. 



196 Inland Fishes of Massachusetts 




Atlantic Needlefish: Coastal records are not 
indicated. 



notes. Needlefishes, particularly the Atlantic 
Needlefish, are sometimes called "gars" or "gar- 
fishes." However, they are not related to the true gars 
(Lepisosteidae) although they superficially resemble the body form of the 
Longnose Gar, Lepisosteus osseus. In the southern parts of their range, At- 
lantic Needlefish are often used for bait for large sport fishes such as mar- 
lins. Needlefishes are eaten in many parts of the world, but not usually in 
the United States, perhaps because of the green color of their bones. 



references. Andrews 1973 (juveniles, Nantucket Harbor); Collette 1968, 
Collette et al. 1984 (relationships, development); Curley et al. 1975 (Bass 
River); Fiske et al. 1967 (Pleasant Bay); Fiske et al. 1968 (Westport River); 
Hardy 1978 (review, development); Hoff and Ibara 1977 (Slocums River). 



Family and Species Accounts 197 



Killifish, Pupfish, and 
Livebearer Families 

Fundulidae, Cyprinodontidae, and Poeciliidae 



Killifishes and pupfishes are closely related families within the order Cy- 
prinodontiformes. This diverse order contains up to 800 species that are 
widely distributed in North America, South America, Europe, Asia, and 
Africa. Until very recently, killifishes (Fundulidae) and pupfishes (Cypri- 
nodontidae) were placed in a single family; however, these two groups are 
now recognized as distinct families. The 48 species of killifishes are endemic 
to North America. Sometimes called minnows, topminnows, bull-minnows, 
or toothed-minnows, these fishes can be distinguished from the true min- 
nows (Cyprinidae) by their toothed jaws. The name killifish most likely 
comes from the Dutch word "kills," meaning small waterways, a favorite 
habitat. Five species of killifishes live in Massachusetts, but one, the Striped 
Killifish, Fundulus majalis, never enters freshwater and is included only in 
the key for reference. The Cyprinodontidae is comprised of about nine gen- 
era and approximately 100 species. Only one species, the Sheepshead Min- 
now, is found in Massachusetts. A number of the pupfishes inhabit the 
deserts of the American Southwest and are often restricted to extremely 
small spring systems. Some pupfishes are able to survive in water that can 
reach at least 110°F. Due primarily to human manipulation of the region's 
water supplies, many pupfish species are threatened, and some are already 
extinct. Killifishes and pupfishes can tolerate low dissolved oxygen and a 
wide range of salinities and temperatures and are able to survive in habitats 
too variable and severe for other types of fishes. The estuarine species, in 
particular, can often survive in small pools that are exposed to high tem- 
peratures and become hypersaline. Killifishes have been extensively stud- 
ied and used as laboratory animals in part because of their ability to survive 
extreme environmental conditions. 

A member of the livebearer family, Poeciliidae, the Mosquitofish, Gam- 
busia affinis, was found for the first time in Massachusetts by Steve Hurley 
(MDFW) in late September 1999. Numerous individuals were found in the 
Quashnet area of Cape Cod. This group of fishes has internal fertilization 
and live young and is related to the guppies of the aquarium trade. The 



198 Inland Fishes of Massachusetts 



Cape Cod population appears to have reproduced in the wild in 1999, but 
we cannot predict if the population will survive New England winters. 

references. Able 1984 (spawning and development); Atz 1986 (labora- 
tory studies); Deacon et al. 1979 (rare and threatened); Griffith 1974 (Fun- 
dulus environmental tolerances); Parenti 1981 (phylogenetic relation- 
ships); Relyea 1983 {Fundulus); Rosen 1973 (review); Wiley 1986 {Fundulus 
relationships). 



Key to Massachusetts Killifishes and Pupfishes 

Note: Mosquitofishes are not included in the key, but they can he easily told from the related 
killifishes by a dark, tear-drop-like mark below the eye and by the dorsal fin that is almost 
entirely (females and young) or entirely (males) behind the anal fin base. The males have 
slender, modified anal fins that serve as their reproductive organ. 



la. Body deep and robust; pectoral fins 
reach a point below dorsal origin; jaw 
teeth wide, with 2 to 3 points. Sheeps- 
head Minnow, Cyprinodon variegatus, 
page 201, Plate 45. 



lb. Body elongate; pectoral fins end well 
before a point below the dorsal fin origin; 
teeth conical. Go to 2. 




Family and Species Accounts 199 



2a. Jaw teeth strongly conical, usually in 
a single row; fewer than 30 lateral line 
scales; scales outlined with fine mark- 
ings; occasional with diffuse bands on 
body. Rainwater Killifish, Lucania parva, 
page 209, Plate 44. 

2b. Jaw teeth pointed and set in bands, 
often buried in tissue; more than 30 lat- 
eral line scales, not outlined as above; 
usually with bands or stripes on body. 
Go to 3. 




3a. Lateral scales usually 40 or more; dis- 
tance from dorsal origin to end of caudal 
peduncle about equal to distance from 
dorsal origin to eye. Banded Killifish, 
Fundulus diaphanus, page 203, Plate 46. 




3b. Lateral scales less than 35; distance 
from dorsal origin to end of caudal pe- 
duncle much less than the distance from 
dorsal origin to eye. Go to 4. 




4a. Snout and upper jaw relatively long 
and pointed (when viewed from side and 
above); prominent black, irregular stripes 
or bars on body. Striped Killifish, Fundu- 
lus majalis. See family account. 





iff} V ■'.'■ "y : % 








d 

9 



4b. Snout and upper jaw relatively short 
and rounded; stripes never present; verti- 
cal bars not prominent or absent. Go to 5. 





200 Inland Fishes of Massachusetts 



5a. Narrow middorsal band from nape to 
dorsal origin; base of dorsal fin shorter 
than base of anal fin; 8 dorsal fin rays. 
Spotfin Killifish, Fundulus luciae, page 
207, Plate 47. 




5b. Middorsal band absent (only a small 
spot at dorsal origin); base of dorsal fin 
equal to or greater than length of the 
base of the anal fin; dorsal fin rays more 
than 11. Mummichog, Fundulus hetero- 
clitus, page 205, Plate 48. 




Sheepshead Minnow 

Cyprinodon variegatus Lacepede 1803 



Native 
plate 45 




identification. Sheepshead Minnows have a stout, deep body and flat- 
tened jaw teeth with two to three cusps (see key Figure la). The end of the 
pectoral fin always reaches a point well behind the origin of the dorsal fin. 
Adult females have a conspicuous dark spot on the dorsal fin while adult 
males have a distinctive dark edge on the caudal fin. Breeding males be- 
come light to steel-blue dorsally and yellow to salmon color ventrally. 

selected counts. D 11-12; A 10-11; Scales 24-27. 

size . Sheepshead Minnows are small fish, rarely reaching 3 inches TL. 
Males are larger than females. 



Family and Species Accounts 20 1 







Sheepshead Minnow: coastal records are not 
indicated. 



natural history. In Massachusetts, Sheeps- 
head Minnows inhabit primarily brackish and marine 
environments. They are frequently found in salt marshes 
and tidal creeks and occasionally enter tidal freshwater. Sheepshead Min- 
nows are omnivorous, eating a wide variety of animal and plant material, 
including small invertebrates, zooplankton, small fishes, and various ma- 
rine algae. They are pugnacious fishes, fighting vigorously with members of 
their own species and with other fishes during an extended midspring to 
late-summer spawning season. Spawning occurs in shallow water, usually 
in association with aquatic vegetation. During spawning, the male holds 
the female while the eggs are released. Eggs mature a few at a time, and 
they are released as they ripen. Laid singly or in small groups throughout 
the spawning season, the eggs sink and stick to aquatic vegetation or other 
substrates. The eggs hatch in less than a week, and juveniles mature in un- 
der a year. 



> m 

i M0 

* 






distribution and abundance. Cape Cod is the northernmost part 
of the Sheepshead's range. In these northern areas they live primarily in 
brackish and marine environments, but in Florida and along the Gulf 
coasts of the United States and Mexico, Sheepshead Minnows often live in 
completely freshwater habitats. In Massachusetts, Sheepshead Minnows 
seldom move beyond the influence of the tide. They are common on the 
south shore of Cape Cod, in areas along the Elizabeth Islands, and on 
Martha's Vineyard and Nantucket. They are occasionally found in tidal 
freshwater or in low salinity coastal overwash ponds. There are no records 
from the Gulf of Maine north of Cape Cod. 



202 Inland Fishes of Massachusetts 



notes. Sheepshead Minnows have been divided into four subspecies with 
C. v. ovinus found in New England. As in the Rainwater Killifish, the New 
England population of Sheepshead Minnow exhibits some noticeable mor- 
phological variations when compared with the southern populations. 

references. Bigelow and Schroeder 1953 (general); Breder and Rosen 
1966 (reproduction); Hardy 1978 (development, review); White et al. 1986 
(life history); Stallsmith 1997 (Nantucket). 



Banded Killifish Native 

Fundulus diaphanus Lesueuv 1817 plate 46 







Itf 






1 - - 



identification. Banded Killifish are more elongate and have a narrower 
caudal peduncle than other Massachusetts killifishes; the distance from the 
dorsal origin to the base of the caudal peduncle usually reaches mideye or 
beyond. Banded Killifish usually have more than 41 lateral scales and have 
18 to 22 dark bands along the sides of the body. In males, the pelvic and 
anal fins are sometimes edged with white, and their body color becomes 
more intense with shades of blue during the spawning season. 

selected counts. D 12-15; A 10-13; Scales 39-49. 

size. Banded Killifish of about 3 inches TL are most commonly encoun- 
tered, but specimens of almost 6 inches TL (117 mm SL) have been col- 
lected from Massachusetts. 

natural history. Banded Killifish typically inhabit freshwater but oc- 
casionally enter slightly brackish water. An adaptable species, they may be 
found in a wide variety of habitats, including ponds and lakes or streams 



Family and Species Accounts 203 




Banded Killifish. 






«t 






and rivers with moderate flow. Areas with clear 
water, some aquatic vegetation, and sandy bot- 
toms, however, appear to be their preferred habitat. 
Banded Killifish, particularly the young, school in shal- 
low water along the shore. Spawning occurs from midspring to midsummer, 
as males establish small territories along the edges of aquatic vegetation. 
Males and females temporarily form pairs, and eggs are laid singly or in 
small groups in the vegetation. Males are active during spawning and vigor- 
ously herd the females to their territories and hold the females close to their 
bodies with their fins. Spawning is probably protracted, and females release 
only a few eggs at a time; the eggs hatch in one to two weeks. Banded Killi- 
fish are primarily carnivorous. Many different aquatic invertebrates are 
taken at various depths, and not just from the surface, as the morphology of 
their upturned mouths might suggest. Fish eggs and larvae are also eaten. 
Banded Killifish are preyed upon by many warm- and cold-water game 
fishes, birds, and mammals. As a defensive behavior, Banded Killifish will 
burrow into the substrate when threatened by a potential predator. 






distribution and abundance. Banded Killifish are common where 
found in Massachusetts. There are records of this species from most of the 
major river drainages; however, populations seem to be somewhat local- 
ized within any given drainage. 

notes. The eastern subspecies of the Banded Killifish, F. d. diaphanus, is 
found from South Carolina north to Newfoundland. It is most closely re- 
lated to the Waccamaw Killifish, F. waccamensis, which is endemic to two 
lakes in North Carolina. 



204 Inland Fishes of Massachusetts 



references. Baker-Dittus 1978 (ecology); Colgan 1974 (defensive be- 
havior); Godin 1986 (schooling); Hardy 1978 (development); Kenney 1981 
(early age and growth, MA); Keast and Webb 1966 (general biology); Scott 
and Crossman 1973 (general biology); Weisberg 1986 (ecology); Wiley 1986 
(relationships). 



Mummichog 

Fundulus heteroclitus (Linnaeus 1766) 



Native 
plate 48 




identification. Mummichogs are stout-bodied, with 11 to 12 dorsal fin 
rays and fewer than 36 lateral scales. They lack the jet black banding and 
longitudinal striping found on Striped Killifish. Banded Killifish are more 
elongate and have more bands than Mummichogs. Juvenile Mummichogs 
might be confused with adults of the locally uncommon Spotfin Killifish 
(see Spotfin Killifish account). Mummichogs are light yellow to olive with 
faint bands or irregular markings on the body. The belly is light in color, 
varying in hue from white to yellow. A dark spot is often found on the pos- 
terior margin of the dorsal fin in males. 

selected counts. D 11-12; A 1 1-12; Scales 34-36; GR 9-11. 

size . Mummichogs are commonly 3 to 4 inches TL, but the maximum size 
is about 6 inches TL. The largest Massachusetts specimen that we have 
measured is about 5 inches TL (111 mm SL). 

natural history. Mummichogs are among the hardiest of fishes and 
can survive in water temperatures of at least 90°F, and salinities of fresh- 
water to well over that of seawater. Mummichogs are most commonly 
found in bays, estuaries, saltmarsh pools, and tidal freshwater. Spawning 



Family and Species Accounts 205 



•L-* 










Mummichog: coastal records are not indicated. 

begins as early as May and continues into the 
early summer, at which time males aggressively 
chase females and rival males. Egg laying is linked to 
photoperiod, water temperature, and tidal cycles. Mum 
michogs use the intertidal marsh for spawning during spring tides. Eggs 
are deposited singly or in small groups near the high water mark. The eggs 
remain out of water for much of their development. In New England, fe- 
males deposit their eggs in sand or algal mats. In other areas, eggs are often 
laid in empty mussel shells or at the base of Spartina leaves. Hatching is 
triggered by the next high tide, when water again completely covers the 
fully developed eggs. Although the eggs are out of water for much of their 
development, surprisingly little egg mortality occurs. The omnivorous 
Mummichog's diet consists of small invertebrates, including amphipods, 
crustaceans, and mollusks. This species also consumes small fishes, includ- 
ing smaller Mummichogs and Mummichog eggs. Plant material and detri- 
tus are frequently ingested but are apparently not nutritionally important. 



■»..- 



V 



4k 



distribution and abund ance . The northern subspecies, F. h. ma- 
crolepidotus, is found north of Long Island, NY, and is abundant all along 
the coast of Massachusetts. This species is most often encountered in 
coastal marsh creeks, ditches, and tide pools. Mummichogs frequently 
enter tidal freshwater and coastal ponds that vary in salinity. 

notes. Mummichogs are often used as laboratory animals, particularly 
in studies offish development, the endocrine system, or pollution. They 
are sometimes called "salt water minnows" or just "minnows" and are fre- 
quently used as bait. This abundant species plays an important role in 



206 Inland Fishes of Massachusetts 



coastal marsh ecology as it affects invertebrate populations and provides 
food for many species of birds and fishes. 

references. Able and Felley 1986 (morphological variations); Atz 1986 
(laboratory use); Collette and Hartel 1988 (Pamet River); Eisler 1986 (pollu- 
tion); Kneib 1984, 1986 (ecology); Relyea 1983; Taylor 1986; Able and Hata 
1984 (reproduction); Weisberg 1986 (competition); White et al. 1986 (life 
history) . 



Spotfin Killifish Native 

Fundulus luciae (Baird 1 855) plate 47 




identification. Spotfin Killifish can be told from the other Fundulus 
by a more posterior dorsal fin that has only eight rays. Spotfin Killifish are 
most similar to young Mummichogs but have a dorsal midline stripe from 
the nape to the dorsal fin origin (see key Figure 5a). 

selected counts. D 8; A 10; Scales 34-36. 

size. Spotfin Killifish are the smallest member of the genus Fundulus. 
Adults rarely reach 2 inches TL. 

natural history. Spotfin Killifish inhabit quiet bays and estuaries. 
They are typically found in the upper regions of salt marshes and on rare 
occasions in tidal freshwater. Their preferred habitat seems to be high in- 
tertidal marshes that do not flood on every tide. In these locations, Spotfin 
Killifish are found in shallow, often temporary pools and have been ob- 
served swimming in areas of marsh vegetation covered by as little as one- 
quarter inch of water. Spawning, which occurs in late spring and summer, 
is most likely associated with the cycle of high tides. Spotfin Killifish have 



Family and Species Accounts 207 



;?' f : 




Spotfin Killifish. 

a maximum life span of two years and are 
sexually mature at two to three months. Their 
diet consists of small invertebrates, which are 
picked from the substrate, fish eggs, and larvae. 



> 



o < -d :S \ 



distribution and abundance. Southern Massachusetts is the north- 
ernmost part of the range of the Spotfin Killifish. The first Massachusetts 
record is based on our collection of seven reproductively active males and 
females (24 to 29 mm SL) that we found in the Palmer River, Rehoboth, on 
June 7, 1980. These specimens were collected in a shallow mosquito ditch 
and over a Spartina marsh flooded to 1 or 2 inches by a high spring tide. In 
June 1999, Bruce Stallsmith found this species common in a similar habitat 
along the Palmer River in Swansea and Rehoboth. 

notes . In the past, this species was considered rare; however, recent stud- 
ies show that it is locally common in some parts of its range. The lack of 
records of Spotfin Killifish is most likely due to the species' preference for 
high tidal marsh habitats that biologists seldom sample for fishes. It is 
also probable that this species is often mistaken for the young of other 
killifishes. 

references. Able et al. 1983 (status, NJ); Brill 1987 (natural history, prop- 
agation); Hardy 1978 (review, development); Kneib 1978 (general biology); 
Kneib 1984 (larval and juvenile ecology); Weisberg 1986 (ecology). 



208 Inland Fishes of Massachusetts 



Color Plates 



The following photographs were taken in Massachusetts by 
K.E. Hartel unless indicated otherwise. All measurements are 
in SL. 




1) American Brook Lamprey, adult 135mm, Blackstone Drainage, 
1981, MCZ 62174. Photo by B. Byrne (MDFW). 




2) Sea Lamprey, transformed juvenile 156mm, Shawsheen River, 
Lawrence, 1998, MCZ 155273. 




3) Shortnose Sturgeon, adult (above), and Atlantic Sturgeon, juvenile 
(below), Merrimack River, 1990. Photo by Boyd Kynard (USFW). 




4) Blueback Herring, adult 192mm, Charles River, 1999, MCZ 
99410. 





"h 










■r 


'kP^^P^S 




; !?w'^ 










..... r . • ■ 










"^ 


3k 


mm Wm 












* 

















5) Blueback Herring, juvenile 61mm (above), MCZ 58160; 
Alewife, juvenile 56mm (below), MCZ 58159. Both Herring River, 
Wellfleet, October 1981. 




6) American Shad, adult. Photo by D. Flescher, courtesy 
American Fisheries Society. 




7) Gizzard Shad, adult 235mm, Connecticut River, North- 
hampton, 1986, MCZ 64569. 




8) Rudd, adult 206mm, Charles River, Boston, 1991, 
MCZ 95616. 




9) Golden Shiner, adult 120mm, Assabet River, 
Concord Drainage, 1988, MCZ 99392. 




10) Goldfish, adult, aquarium specimen. 




11) Common Carp, juvenile 54mm, Connecticut 
River, 1980, MCZ 57115. 




12) Northern Redbelly Dace, adult 35mm, Deerfield 
Drainage, 1979, MCZ 54699. 




13) Lake Chub, adult 77mm, Middle Branch Westfield River, 
1950, MCZ 54694. 




14) Eastern Silvery Minnow, adult 71mm, Connecticut River, 
Hadley, 1950, UMA 35-1. 




15) Spottail Shiner, adult 77mm, Charles River, 1988, MCZ 99409. 




16) Mimic Shiner, adult 49mm, Connecticut River, 1980, MCZ 
57159. 




17) Common Shiner, adult 57mm, Deerfield Drainage, 1981, 
MCZ 56516. 




18) Fallfish, adult 146mm, Center Brook, Blackstone 
Drainage, 1988, MCZ 99404. 




19) Creek Chub, adult 69mm, South River, Conway, 1980, MCZ 
57263. 




20) Bridle Shiner, adult 36mm, Parker Drainage, 1988, MCZ 
95249. 




21) Blacknose Dace, adult 61mm, S. Wachusetts Brook, 
Nashua Drainage, 1988, MCZ 99395. 



22) Longnose Dace, adult 94mm, Gates Brook, Nashua 
Drainage, MCZ 99398. 




23) Bluntnose Minnow, adult 64mm, Housatonic River, 1979, MCZ 
56433. 




24) Fathead Minnow, adult 42mm, Housatonic River, 1979, MCZ 
56431. 




25) Creek Chubsucker, subadult 55mm, Merrimack 
Drainage, 1988, MCZ 99412. 




26) White Sucker, adult 180mm, Center Brook, Blackstone 
Drainage, 1988, MCZ 99397. 




27) Longnose Sucker, adult 172mm, North River, Deerfield 
Drainage, 1981, MCZ 57101. 




28) White Sucker, juvenile (above) and Longnose Sucker, juvenile 
(below). Each about 40mm, both Housatonic Drainage, 1979. 




29) Central Mudminnow, adult 36mm, Connecticut Drainage, 
1980, MCZ 56943. 




30) Trout-perch, adult 72mm, Green River, Housatonic Drainage, 
1940, MCZ 54922. 




31) White Catfish, adult 255mm, Charles River, 1988, MCZ 
79899. 




32) Brown Bullhead, adult 160mm, Center Brook, 
Blackstone Drainage, 1988, MCZ 99399. 




33) Yellow Bullhead, adult 95mm, Merrimack Drainage, 
1988, MCZ 99413. 




34) Channel Catfish, used by permission of R.S. Wydoski 
and R.R. Whitney, 1979. Inland Fishes ofWashington, 
University ofWashington Press, Seattle. 




35) Tadpole Madtom, adult 78mm, Howe Pond, Chicopee 
Drainage, 1978, MCZ 54224. 




36) Redfin Pickerel, adult 127mm, Center Brook, 
Blackstone Drainage, 1988, MCZ 99400. 




37) Chain Pickerel, adult 191mm, Center Brook, 
Blackstone Drainage, 1988, MCZ 99402. 




38) Atlantic Tomcod, juvenile 60mm, Taunton River, 1980, 
MCZ 57295. 




39) Burbot, juvenile, Third Connecticut Lake, NH, 1985, 
MCZ 63017. 




40) Atlantic Salmon, juvenile 145mm, Gates Brook, Nashua 
Drainage, 1988, MCZ 99393. 




41) Brown Trout, adult 170mm, Gates Brook, Nashua 
Drainage, 1988, MCZ 99406. 




42) Brook Trout, adult 150mm, S. Wachusetts Brook, Nashua 
Drainage, 1988, MCZ 99407. 




43) Rainbow Smelt, adult 101 mm, Maiden Brook, Nashua 
Drainage, 1984, MCZ 93565. 





44) Rainwater Killifish, adult 30mm, Buzzards Bay Drainage, 
1980, MCZ 57136. 




45) Sheepshead Minnow, adult 36mm, Cape Cod Drainage, 
1980, MCZ 57142. 




46) Banded Killifish, adult 70mm, Charles River, Boston, 
1980, MCZ 59059. 





47) Spotfin Killifish, adults, female 27mm (above), male 
24mm (below), Palmer River, Rehoboth, 1980, MCZ 57600. 




48) Mummichog, adult 60mm, Neponset River, 1988, 
MCZ 99421. 





49) Fourspine Stickleback, adult 32mm, Neponset River, 1980, MCZ 
99420. 




50) Threespine Stickleback, adult 35mm, Olmstead Park, Boston, 
1988, MCZ 99417. 




51) Blackspotted Stickleback, adult 37mm, Cohasset, 1977, MCZ 
52753. 




52) Ninespine Stickleback, adult 47mm, Eel River, Cape Cod 
Drainage, 1988, MCZ 99418. 




53) Inland Silverside, adult 45mm (above), Charles River, 1979, 
MCZ 56124; Atlantic Silverside, adult 51mm (below), Neponset 
River, 1979, MCZ 56238. 




54) Slimy Sculpin, adult 70mm, S. Wachusetts Brook, Nashua 
Drainage, 1988, MCZ 99405. 







55) Hogchoker, subadult 57mm, Palmer River, Rehoboth, 
1981, MCZ 58244. 




56) White Perch, adult 130mm, Charles River, 1988, MCZ 
99411. 




57) Banded Sunfish, adult, Merrimack 
Drainage, 1988. 




58) Black Crappie, juvenile 79mm, Charles 
River, 1979, MCZ 56551. 




59) Bluegill, adult 143mm, Charles River, 1988, 
MCZ 99422. 




60) Rock Bass, 120mm, Merrimack River, New 
Hampshire state line, 1990, MCZ 95848. 




61) Redbreast Sunfish, adult 110mm, Charles 
River, 1988, MCZ 99426. 




62) Pumpkinseed, adult 95mm, Assabet River, 
Concord Drainage, 1988, MCZ 99401. 




63) Smallmouth Bass, subadult, 169mm, Johns 
Creek, Virginia, 1984. Photo by R.E. Jenkins. 




64) Largemouth Bass, adult 115mm, Assabet 
River, Concord Drainage, 1988, MCZ 99394. 




65) Largemouth Bass, juvenile 60mm (above), Merrimack 
River, 1979, MCZ 57279; Smallmouth Bass, juvenile 56mm 
(below), Connecticut River, 1980, MCZ 57344. 




66) Yellow Perch, adult 115mm, Charles River, 1988, MCZ 99419. 




67) Swamp Darter, adult 29mm, Gibbs Pond, Nantucket, MCZ 
58231. 




68) Tessellated Darter, adult 60mm, Center Brook, Blackstone 
Drainage, 1988, MCZ 99403. 



Rainwater Killifish 

Lucania parva (Baird and Girard 1855) 



Native 
plate 44 




identification. Rainwater Killifish have caninelike teeth in a single 
row, relatively short bodies with large scales, and pectoral fins that extend 
back to a point just before the origin of the dorsal fin. Each scale is deli- 
cately outlined to give the body a crosshatched appearance. Most males 
have a small, dark spot on the lower anterior edge of the dorsal fin; during 
breeding the fins have an orange-red wash, and the dorsal, caudal, anal, 
and pelvic fins have dark edging. 

selected counts. D 11-12; A 10-11; Scales 25-26. 

size. Rainwater Killifish are small, usually reaching only 1.5 inches TL. The 
largest Massachusetts specimen that we have seen is about 2 inches TL. 

natural history. This small fish is tolerant of a wide range of water 
conditions, from freshwater to hypersaline pools in salt marshes. A school- 
ing species, generally associated with aquatic vegetation, Rainwater Killi- 
fish are most often found in salt marsh creeks and estuaries. In some areas, 
however, this species naturally inhabits large rivers and streams. In Massa- 
chusetts, they are most common in coastal marshes, creeks, and overwash 
ponds, although they occasionally move into tidal freshwaters. Rainwater 
Killifish travel in schools and may move from brackish to freshwater during 
the breeding season. Males become territorial during the late spring and 
summer spawning season, defending small areas near aquatic vegetation. 
Females are actively courted and enticed into the males' territories by a 
series of energetic displays. If receptive, a female follows a male to his ter- 
ritory and deposits several eggs close to aquatic vegetation, or other appro- 



Family and Species Accounts 209 




Rainwater Killifish. 

priate structures, and the eggs are then quickly 
fertilized by the male. Eggs hatch in 6 to 14 days, 
depending on water temperature. The diet of Rain- 
water Killifishes is comprised of a variety of small aquatic 
invertebrates, including mosquito larvae and crustaceans. 












\MJL 



.<*■ 



distribution and abundance. In Massachusetts, Rainwater Killi- 
fish are common in many of the drainages to Nantucket Sound and to Buz- 
zards and Narragansett bays. They are common in the tidal freshwaters of 
Martha's Vineyard, where they are also found in a freshwater pond at Felix 
Neck. Rainwater Killifish seem to be rare on Nantucket, where the first 
records were brought to our attention by B. Stallsmith in 1995. 

notes. Hubbs and Miller (1965:35-36) state that "...the most trenchantly 
distinct of the local forms [of L. parva] inhabits southern New England" and 
"...Were it not for the irregularity in the clines, this New England race would 
warrant separation as the nominate subspecies." They point out that simi- 
lar patterns of differentiation have been noted for southern New England 
populations of Sheepshead Minnow, silversides, and Hogchoker. 

references. Hardy 1978 (development, review); Hubbs and Miller 1965 
(description, distribution, systematics, variation); Moyle 1976 (general biol- 
ogy); Beck and Massmann 1951 (movement); Stallsmith 1997 (Nantucket). 



210 Inland Fishes of Massachusetts 



Silverside Family 

Atherinopsidae 



The silverside family, formerly called the Atherinidae, consists of small 
fishes, usually less than 8 inches in total length, that are widely distributed 
throughout tropical and temperate regions of the world. They are related to 
the killifishes and pupfishes (Fundulidae and Cyprinodontidae). Silversides 
inhabit marine, brackish, and freshwaters. Some species inhabit Andean 
lakes, and one species, the Brook Silverside, Labidesthes sicculus, is com- 
mon in freshwaters of the central and southeastern United States with its 
northeastern limit in New York. As their common name indicates, 
atherinopsids are silvery, and somewhat translucent, with a distinct, silver 
midlateral band. They have two dorsal fins; the first is small, inconspicu- 
ous, and separated from the second. The lateral line is reduced or absent, 
and the pelvic fins are located approximately at midbody. Silversides have 
small mouths with fine teeth. They often school in large numbers. In many 
parts of the world, silversides are fried whole or dried for human consump- 
tion and are often called "whitebait." 

references. Bigelow and Schroeder 1953 (MA); Chernoff et al. 1981, 
Dyer and Chernoff 1996 (systematics); Gosline 1948 (speciation); Johnson 
1975, Parenti 1993 (relationships). 



Key to Massachusetts Silversides 



la. Predorsal scales fewer than 16; anal 
fin rays usually fewer than 18. Inland Sil- 
verside, Menidia beryllina, page 212, 
Plate 53. 




Family and Species Accounts 211 



lb. Predorsal scales usually more than 
20; anal fin rays usually more than 22. 
Atlantic Silverside, Menidia menidia, 
page 214, Plate 53. 




Inland Silverside 

Menidia beryllina (Cope 1866) 



Native 

PLATE 53 




identification. Silversides are silvery fishes with two dorsal fins and 
one weak anal spine. Inland Silversides are similar to Atlantic Silversides 
but have shorter anal fins and larger scales. Inland Silversides have fewer 
than 16 predorsal scales and 15 to 16 (rarely 20) anal rays. They are a light 
silver green to waxen yellow dorsally with a well-defined midlateral silver 
stripe. The dorsal and caudal fins are occasionally washed with orange. 

selected counts. D IV-VII, 8-10; A 1,13-19; Scales 36-42. 

size . Inland Silversides are smaller than the Atlantic Silversides and rarely 
reach 4 inches TL. The largest Massachusetts specimens that we have ex- 
amined measure just over 3 inches TL (67 mm SL). 

natural history. In Massachusetts, Inland Silversides are frequently 
found in bays, salt marshes, estuaries, coastal freshwater, and overwash 
ponds. They commonly enter areas of freshwater above the influence of the 
tides but rarely travel far upstream. Inland Silversides are often found with 
Atlantic Silversides and usually outnumber them in less saline areas. They 
are commonly found in large schools that remain in the estuaries most of 



212 Inland Fishes of Massachusetts 




Inland Silverside: coastal records are not indicated. 

the year and have relatively small home ranges. 
Spawning occurs in the summer when schools of 
silversid es concentrate in shallow water. The sticky 
eggs are often laid in the vegetation of the intertidal zone 
at high tide. These eggs quickly sink and adhere to the vegetation or sandy 
bottom. The eggs hatch in one to two weeks, depending on water tempera- 
ture. The juveniles become mature at one year. Diet is varied and a wide 
range of plant and animal material is eaten, including zooplankton, shrimp, 
amphipods, mollusks, worms, larval fishes, fish eggs, and, to a lesser extent, 
algae and detritus. 



distribution and abundance. In Massachusetts, the Inland Silver- 
side is most commonly found in the coastal streams, bays, and estuaries on 
the south side of Cape Cod. North of Pleasant Bay on the outer arm of Cape 
Cod, Inland Silversides are much less abundant but may be expected in 
most of the bays and estuaries south of Nahant. Bigelow and Schroeder 
(1953) found only one record from north of the Cape, but we found that In- 
land Silversides were common to abundant in the Massachusetts Bay area, 
including the Charles River in Cambridge and Boston during the mid- to 
late 1970s. However, recent surveys (1983-1997) in the Charles River have 
not found this species. The abundance north of Cape Cod may be cyclic, 
expanding or contracting depending on environmental factors. 

notes. This species has also been variously called the "waxen" or "tide- 
water" silverside. A southern New England subspecies, M. b. cerea, has 
been described, but its status has not been recently reviewed. 



Family and Species Accounts 213 



references. Bengtson 1985, Middaugh et al. 1986 (reproduction); Bige- 
low and Schroeder 1953 (general biology); Collette and Hartel 1988 (Massa- 
chusetts Bay); Kendall 1902, Johnson 1975, Chernoff et al. 1981 (systemat- 
ics); Korth and Fitzsimons 1987 (karyotype); Hoff 1972 (movement, MA). 



Atlantic Silverside Native 

Menidia menidia (Linnaeus 1766) plate 53 




identification. Silversides are silvery fishes with two dorsal fins and 
one weak anal spine. Atlantic Silversides are similar to Inland Silversides 
but have longer anal fins and smaller scales. Atlantic Silversides usually 
have more than 20 crowded predorsal scales and 22 to 25 anal rays. They 
are a silver-green dorsally and have a well-defined midlateral silver stripe. 

selected counts. D III-VII,7-1 1; A 1,19-29; Scales 43-55. 

size. Atlantic Silversides commonly reach 4.5 inches TL, and individuals 
up to 5.5 inches TL have been recorded. 

natural history. Atlantic Silversides are inshore marine fishes that are 
frequently found in bays, salt marshes, and estuaries but only rarely enter 
areas of freshwater above the influence of the tides. They are commonly 
found in large schools in areas of sandy bottoms and aquatic vegetation. 
Silversides often congregate in shallows, particularly when high tides have 
partially submerged the shore vegetation. In winter, most silversides move 
into deeper water, but some may survive in shallow water under the ice. 
Winter mortality is occasionally high as large winter die-offs have been re- 
ported. In Massachusetts, protracted spawning occurs from mid-May to 
late July and eggs are often laid in the vegetation of the intertidal zone, usu- 
ally at high tide. The eggs, which have tufts of filaments, sink and adhere to 



214 Inland Fishes of Massachusetts 




•♦<. 






Atlantic Silverside: coastal records are not indicated. 




• '*» 






> V 

> -+J; 



the vegetation or sandy bottom. Virtually all of 
the eggs are laid above the mean low tide level and 
are exposed to the air for parts of their development. a3»»a *i 

Eggs are sometimes found in large numbers, occasionally 
forming large sheets and clusters. The eggs hatch in one to two weeks, de- 
pending on water temperature, and the young mature at one year. Most 
Atlantic Silversides in Massachusetts die before they reach two years of age 
Diet is varied; Atlantic Silversides eat a wide range of plant and animal 
material. 






A 



distribution and abundance . Atlantic Silversides are found along 
the Atlantic coast of North America from the Gulf of Saint Lawrence to 
northern Florida. In Massachusetts, this species is common to abundant 
along virtually the entire coast. However, the abundance of Atlantic Silver- 
sides may be cyclic and is also linked to the availability and health of salt 
marshes. 

notes. Like the Inland Silverside, the Atlantic Silverside is an important 
forage fish and is readily eaten by game fishes, such as the Bluefish, Poma- 
tomus saltatrix, and the Striped Bass. It is also heavily preyed upon by birds, 
particularly terns and herons, when the silversides congregate in shallow 
water. Atlantic Silversides are a popular, but soft, bait fish. 

references. Bigelow and Schroeder 1953, Clayton et al. 1978 (review, 
MA); Bengtson et al. 1987 (reproduction); Conover 1979, 1982, Conover and 
Ross 1982 (biology, MA); Middaugh 1981 (reproductive ecology). 



Family and Species Accounts 215 



Mullet Family 

Mugilidae 



Mullets are primarily marine fishes that are found worldwide in inshore 
tropical and temperate waters. Many species regularly enter rivers and 
streams, and some spend their entire lives in freshwater. Fishes of this 
family often form loose schools and feed in bays and estuaries. They have 
a specialized pharyngeal organ and a long, coiled gut that allows them to 
process their food. Diet is variable, but plant material and detritus are often 
ingested. In some regions, mullets are commercially important fishes. 
Recently the relationships of the mullets have been studied in depth, 
and although there is some disagreement all reviewers agree that they 
are closely related to the atherinoids. Of the 70 or so species, only two are 
found as far north as New England. One, the Striped Mullet, frequently en- 
ters the freshwaters. The other species, the White Mullet, Mugil curema, is 
similar but seldom enters freshwater. The two species can be separated by 
the following key. 

references. Stiassny 1993, Nelson 1994 (relationships); Harrison and 
Howes 1991 (pharyngeal organ). 



Key to Massachusetts Mullets 

la. Anal fin with 11 elements, 3 spines 
and 8 rays in adults, 2 spines and 9 rays 
in juveniles less than 40 mm SL; scales 
usually absent from second dorsal and 
anal fins (not illustrated). Striped Mullet, 
Mugil cephalus, page 217. 

lb. Anal fin with 12 elements, 3 spines 
and 9 rays in adults, 2 spines and 10 rays 
in juveniles less than 40 mm SL; second 
dorsal and anal fins with rows of scales 
(not illustrated). White Mullet, Mugil 
curema. See comments under Striped 
Mullet. 



216 Inland Fishes of Massachusetts 



Striped Mullet Native 

Mugil cephalus (Linnaeus 1758) 




identification. Juvenile mullets are similar in appearance to adult 
silversides (Atherinopsidae). The anal fin of the Striped Mullet has many 
fewer rays than those of silversides; there are fewer than 9 in mullets and 
more than 12 in silversides. The origin of the anal fin is directly below or 
behind the origin of the second dorsal fin in the mullet, while it is anterior 
to the origin of the dorsal fin in silversides. The White Mullet can be distin- 
guished from the Striped Mullet by the characters in the key (anal counts, 
dorsal and anal fin scales) and the fact that fresh adult Striped Mullet have 
stripes along the body. The sides and belly of the Striped Mullet are silvery 
white, and the upper surfaces of the body a darker blue grey. The young are 
often bright silvery. 

selected counts . D IV, 1,8; A III— 8 

size. Striped Mullet are usually around 20 inches long (450 to 550 mm SL), 
though specimens as large as 2 feet have been recorded (622 mm SL). Males 
are usually slightly smaller than females. 

natural history. Striped Mullet are an inshore marine species, and 
small schools are frequently found in bays, estuaries, and coastal streams. 
Diet consists of small invertebrates, zooplankton, and detritus. Mouthfuls 
of substrate are scooped from the bottom and are handled with the pharyn- 
geal organ and a specialized gizzard. As is typical with many detritivores, 
food is processed in a long coiled intestine. Mullets are nocturnal spawners 
in areas up to 40 to 50 miles offshore and have pelagic larvae and postlarvae 
that return to coastal waters. Spawning is probably uncommon in waters as 



Family and Species Accounts 217 




Striped Mullet: coastal records are not indicated. 



im 

far north as Massachusetts. Most Striped Mullet \ V J r 
found in the waters of this state are the result of 
northerly transport of the pelagic juveniles by ocean 
currents. Mullet are important food for birds and other 
fishes. 




distribution and abundance. The Striped Mullet has a cosmopoli- 
tan distribution and is found in virtually all the temperate and tropical 
inshore areas of the world. In the western North Atlantic, this species has 
been recorded as far north as Nova Scotia but is not usually common north 
of Cape Cod. Indeed, most of the previous identifications of Striped Mullet 
north of Cape Cod are thought to be White Mullet (Scott and Scott 1988). 
However, several hundred Striped Mullet were reported in the vicinity of 
the Pilgrim Nuclear Generating Station and in other areas of Massachusetts 
Bay during November and December 1975. In Massachusetts, Striped Mul- 
let are frequently encountered during the summer months in bays, estuar- 
ies, and freshwater coastal rivers and streams draining the south side of 
Cape Cod and the Islands. 

notes. Striped Mullet are raised in brackish ponds for human consump- 
tion in many parts of the world and have been extensively studied. 

references. Bigelow and Schroeder 1953, Scott and Scott 1988 (old and 
new records); Fairbanks and Lawton 1977 (Mass. Bay records); Stiassny 
1993 (relationships); Thompson 1963, 1997 (biology, nomenclature). 



218 Inland Fishes of Massachusetts 



Stickleback Family 

Gasterosteidae 



Sticklebacks are found in temperate marine and inland waters of the North- 
ern Hemisphere. The family has about five genera and nine species. Stickle- 
backs show so many inter- and intra-populational differences in morphol- 
ogy and behavior that just one species has been described as more than 
40 different species. Male sticklebacks are brightly colored during breeding 
season and build a nest out of aquatic vegetation using kidney secretions. 
They actively court females, enticing them to the nests with a series of 
highly complex and species-specific displays. Each nest may contain the 
eggs of several different females. Males aggressively guard the nests and 
care for the eggs and newly hatched young. Sticklebacks have been well 
studied in relation to evolution, ecology, ethology, physiology, reproduc- 
tion, and endocrinology. 

references. McLennan et al. 1988 (behavior/relationships); Wootton 
1976 (summary of family); Fitzgerald 1983 (reproduction /behavior); Tin- 
bergen 1952 (behavior); Wootton 1984 (general biology). 



Key to Massachusetts Sticklebacks 



la. Usually 8 or more short, alternately 
inclined dorsal spines, body elongate. 
Ninespine Stickleback, Pungitius pungi- 
tius, page 227, Plate 52. 



lb. Less than 8 medium to long dorsal 
spines, body moderately deep. Go to 2. 



Family and Species Accounts 219 



2a. Pelvic skeleton shows externally as a 
pair of ventro-lateral keels. Fourspine 
Stickleback, Apeltes quadracus, page 221, 
Plate 49. 




2b. Pelvic skeleton with a single median 
posterior extension. Go to 3. 




3a. Pelvic fin with 1 small soft ray; pelvic 
spine lacking ventral cusp (ventral view 
shown); usually with bony lateral keel on 
caudal peduncle; always lacks spots on 
flanks. Threespine Stickleback, Gasteros- 
teus aculeatus, page 223, Plate 50. 




3b. Pelvic fin usually with 2 small soft 
rays; pelvic spine with well-developed 
ventral cusp (ventral view shown); always 
lacks bony lateral keel on caudal pe- 
duncle; dark spots usually present on 
flanks. Blackspotted Stickleback, Gas- 
terosteus wheatlandi, page 225, Plate 51. 




220 Inland Fishes of Massachusetts 



Fourspine Stickleback Native 

Apeltes quadracus (Mitchill 1815) plate 49 




identification. Fourspine Sticklebacks have two ventro-lateral pro- 
cesses of the pelvic girdle (see key Figure 2a) and usually have four or five 
dorsal spines. They are olive-brown dorsally with dark irregular markings, 
and the belly and breast are white to silver in color. Males become much 
darker and the pelvic fins turn bright red during the breeding season. 

selected counts. D IV-V,1 1-12; A 1,9-1 1; Pel 1,2. 

size . The Fourspine Stickleback is a small fish with adults rarely reaching 
2.5 inches TL. 

natural history. Fourspine Sticklebacks are found primarily in salt 
marshes and tidal creeks. They seasonally enter freshwater and may be 
found considerable distances upstream. Fourspine Sticklebacks move into 
fresh and brackish waters from May to July, and the males build small nests 
out of aquatic vegetation. These nests are constructed well above the bot- 
tom by gluing pieces of aquatic vegetation and detritus with a special kid- 
ney secretion. Males actively court any female that swims by the partially 
completed nest. If the female is receptive, she burrows into the top of the 
nest and deposits adhesive eggs. Each female lays approximately 35 eggs; 
however, the number varies with the age and size of the female. After fertil- 
izing the eggs, the male chases the female away and frequently builds sev- 
eral more nests on top of the original nest. The males tend and defend the 
eggs and young. In most populations, the males live only one year, while 
females may live up to three years. Diet includes a wide variety of small 
aquatic invertebrates, particularly amphipods, isopods, dipterans, and zoo- 
plankton as well as aquatic vegetation. 



Family and Species Accounts 22 1 




Fourspine Stickleback: coastal records are not indi- 
cated. 



V" 









\ /- 



I* 






distribution and abundance. In Massa- 
chusetts, Fourspine Sticklebacks are found in estu- 
aries along the entire coast and are common in some 
freshwater streams and ponds in the Cape Cod, Buzzards Bay, and Narra- 
ganset Bay drainages, and on Martha's Vineyard. They are found as far in- 
land as Haverhill on the Merrimack River. 






notes. The Fourspine Stickleback has been observed removing and eating 
external parasites from the Rainwater Killifish. This behavior is called a 
cleaning symbiosis and has been noted in a number offish species. 

references. Bigelow and Schroeder 1953 (general); Schwartz 1965 (life 
history); Baker 1971 (habitat preference); Krueger 1961, Blouw and Hagen 
1987 (morphological variation); Reisman 1963 (reproduction); Rowland 
1974a, 1974b (reproduction); Tyler 1963 (cleaning symbiosis); Wootton 
1976, 1984 (reviews). 



222 Inland Fishes of Massachusetts 



Threespine Stickleback Native 

Gasterosteus aculeatus Linnaeus 1758 plate 50 




identification. Similar to Blackspotted Sticklebacks, Threespine 
Sticklebacks have only one small pelvic ray and lack well-developed ventral 
cusp on the pelvic spine (see key Figure 3a). A lateral keel on the caudal pe- 
duncle is typically present in the Threespine Stickleback but may be absent 
in inland populations. Threespine Sticklebacks are drably colored, varying 
from a silver yellow to light brown and green except during the breeding 
season, when the overall color of the male darkens and the breast and belly 
become a bright red with the eyes turning vivid light blue. 

selected counts. D 111,9-11; A 1,9-11; Pel 1,1. 

size . Marine Threespine Sticklebacks commonly reach 2 to 3 inches TL. 
Individuals of Massachusetts' only known landlocked population are 
smaller, typically ranging from 1 to 1.5 inches TL. 

natural history. In Massachusetts, Threespine Sticklebacks typically 
occur in marine environments and move toward freshwater during the 
spring and early summer spawning season. They are commonly encoun- 
tered in estuaries, salt marshes, and the tidal portions of rivers and streams. 
At the beginning of the spawning season, males construct small, barrel- 
shaped nests of vegetation and detritus on the bottom. Females are at- 
tracted to the nests by an elaborate courtship display known as the "zigzag" 
dance. Males may mate with several females, and individual nests may con- 
tain up to 600 eggs. Care of the eggs and young is left to the males. 

The life span of the Threespine Stickleback is variable between popula- 
tions; some individuals live only a single year, and others up to 3.5 years. 
They become mature at one to three years of age. Threespine Sticklebacks 



Family and Species Accounts 223 




- <t_b£Z /-ii-ULfl ii (™i v i^- 3 * t \ \\S] \y^c £"-> ; 

IAS Va ( K " N y 



Threespine Stickleback: open circles indicate his- l { (yj '\/'J% m Uk i 

toric records where the species may no longer occur; ^\jj i \ > ) V £ff/ 
coastal records are not indicated. 7 Y / ^ "" 



/ "'- 








!<£,. 


l : \ 


:r*> 




• 


<, 


*a$) 



s *y 



eat a variety of small aquatic invertebrates, includ- > ^ /£ s;i 

ing copepods, cladocerans, hemipterans, dipterans, and 
ostracods. Aquatic plants, larval fishes, fish eggs, and mollusks are also 
frequently eaten. However, the specific diet is highly dependent on the 
type of habitat in which the stickleback is living and the seasonal abun- 
dance of prey. 



■s\ 



distribution and abundance. In Massachusetts, Threespine Stickle- 
backs are found along the entire coast in estuaries, salt marshes, and tidal 
creeks. The Threespine Stickleback is often abundant in coastal waters of 
low salinity during the spring and summer spawning season. The only land- 
locked population is located in Boston's Olmsted Park. 

notes. The landlocked population of Threespine Sticklebacks in Olmsted 
Park (which forms the boundary between Boston and Brookline) is unique 
for several reasons. The population has three distinct lateral-plate morphs, 
low numbers of plates without a caudal keel, intermediate numbers of 
plates, or is completely plated. Most Olmsted Park individuals are the com- 
plete or low lateral-plate morphs. In eastern North America, this population 
represents only the fourth record of low plate individuals and is the south- 
ernmost completely freshwater population. This population has been iso- 
lated from the Charles River Basin estuarine population only since the late 
1800s. Within their small range (225-square-yard pool and outflow), these 
animals are common but are susceptible to extinction due to their urban 
habitat. The population is currently listed as Threatened by the Massachu- 
setts Division of Fisheries and Wildlife. It is possible that this population 



224 Inland Fishes of Massachusetts 



was introduced to the pools because the area was planned to be part of 
a natural history museum complex by Frederick Law Olmsted (never- 
completed), and he had designated one of the pools to exhibit sticklebacks. 

references. Bell and Foster 1994, Wootton 1976, 1984 (biology, review); 
Bell and Baumgartner 1984 (Boston, freshwater population); Coad 1983 
(Canada, freshwater populations); Coad and Power 1973 (ecology). 



Blackspotted Stickleback 

Gasterosteus wheatlandi Putnam 1867 



Native 
plate 51 




identification. Blackspotted and Threespine Sticklebacks are similar, 
but Blackspotted Sticklebacks usually have two small pelvic rays and always 
lack the lateral keel on the caudal peduncle. They have well-developed ven- 
tral cusps at the base of the pelvic fin spine (see key Figure 3b). Blackspotted 
Sticklebacks are usually light silver-white with only a faint hint of black 
spots along the sides of the body. During spawning, the body and unpaired 
fins become yellow-green, and the dark spots on the sides of the body and 
head are obvious. 

selected counts. D III, 9-10; A I, 6-8; Pel 1,1-3. 

size. The Blackspotted Stickleback is a small species; individuals over 
2 inches TL are seldom observed. 

natural history. Blackspotted Sticklebacks are the most marine of the 
Massachusetts sticklebacks. They are found from estuaries to well offshore. 
They occasionally move upstream but rarely so far as to enter freshwater. 
Blackspotted Sticklebacks normally remain in brackish environments, such 



Family and Species Accounts 225 







Blackspotted Stickleback: open circles indicate his 
toric records where the species no longer occurs. 



T,.W i ) it..' s 

l u(U 

as salt marshes and bays. Spawning occurs from W . ' . ^ 
late spring to early summer. During this period, Black- >c /tw^ ; . 
spotted Sticklebacks spawn in a manner similar to Three- 
spine Sticklebacks. Variations in their reproductive dance sequences serve 
to isolate these species and prevent hybridization. Blackspotted Stickle- 
backs apparently spawn at one year, and most die shortly thereafter. Fe- 
males have been found to carry approximately 70 to 160 eggs. The diet of 
this species is poorly known but probably consists of aquatic invertebrates 
and juvenile fishes. 



\ \ 



t 






« 4- 



distribution and abundance. The Blackspotted Stickleback has the 
smallest range of any of the American stickleback species. It is found only 
from Newfoundland south to New York. In Massachusetts, the Blackspotted 
Stickleback lives along the entire coast. Freshwater populations are un- 
known in Massachusetts; however, these fish are abundant in areas of low 
salinity in Sesachacha Pond, Nantucket. 

notes. This species was described by Harvard University's anthropologist 
and natural historian, F.W. Putnam, from specimens collected at Nahant, 
Massachusetts. The species is named after R.H. Wheatland, one of the early 
naturalists at the Essex Institute in Salem. 

references. Bigelow and Schroeder 1953, Scott and Crossman 1973 
(general); Coad and Power 1973 (ecology); Reisman 1968, Mclnerney 1969 
(reproduction); Sargent et al. 1984 (plate variations); Wootton 1976 (general 
biology). 



226 Inland Fishes of Massachusetts 



Ninespine Stickleback 

Pungitius pungitius (Linnaeus 1758) 



Native 
plate 52 




identification. Ninespine Sticklebacks have an elongate body and usu- 
ally 8 to 1 1 short dorsal spines that fold alternately from one side to the 
other when collapsed. The upper body of the Ninespine Stickleback is a 
light olive-green with irregular dark markings. The belly and breast are a 
lighter silvery-yellow to white. Males darken considerably during breeding 
season, and their pelvic fin spines turn white. 

selected counts. D VIII-XI(VI-XIII),9-11; A 1,8-10; Pel 1,1. 

size. Ninespine Sticklebacks are generally small fishes; in our recent col- 
lections we have seldom seen specimens over 2 inches TL. However, in a 
series of MCZ specimens collected between 1856 and 1859 from Salem Mill 
Pond, the adults range up to 3.5 inches TL (71 mm SL). 

natural history. In Massachusetts, Ninespine Sticklebacks are found 
in salt marshes and coastal streams. They are often associated with well- 
vegetated freshwater or slightly brackish areas, particularly during the late 
spring to summer spawning season. Small, tunnel-shaped nests are con- 
structed by males from aquatic vegetation and are usually built well above 
the bottom. Females are attracted to the nests by a series of courtship dis- 
plays. If these displays are successful, the female will enter the nest and de- 
posit 30 to 40 eggs. Gravid females contain up to 140 eggs. The eggs in the 
nest are quickly fertilized by the male, and the female is chased away. Males 
may mate with up to seven females in each nest and may construct several 
nests. Males tend the eggs and young until the young are 0.25 to 0.5 inches 
long. Individuals of this species become mature at one year and may live to 
be 3.5 years. Ninespine Sticklebacks are known to eat a wide variety of 



Family and Species Accounts 227 




Ninespine Stickleback: open circles indicate his- 
toric records where the species may no longer occur; 
coastal records are not indicated. 



small aquatic invertebrates and fish eggs and fry, 
even those of their own species. 



distribution and abundance. Massachusetts populations of the 
Ninespine Stickleback are often found in estuaries and salt marshes. 
Coastal freshwater populations seem to be more common in the streams 
of the Southern New England Drainage Area. There is at least one com- 
pletely landlocked population on Martha's Vineyard. Ninespine Stickle- 
backs are often numerous, but they are found far less frequently than the 
other sticklebacks. 

notes. The relationships of the many forms of the Ninespine Stickleback 
in North America and Eurasia are still in question. Like other sticklebacks, 
this species is polymorphic in lateral plate numbers and placement. In the 
Ninespine Stickleback, the lateral plates can be complete, partial, or found 
only on the caudal keel. Haglund et al. (1992) suggest that the name Pungi- 
tius occidentalis (Cuvier) be applied to the North American population. 

references. Griswold and Smith 1973 (life history); Fitzgerald 1983 
(ecology and behavior) ; Ayvazian and Krueger 1992 (lateral plates); Woot- 
ton 1976 (general biology); Haglund et al. 1992 (relationships). 



228 Inland Fishes of Massachusetts 



Pipefish and Seahorse Family 

Syngnathidae 



The seahorse and pipefish family is primarily marine, but a number of spe- 
cies live in freshwater and many others occasionally enter freshwater 
streams. Seahorses and pipefishes have a worldwide distribution in tropical 
and temperate waters. There are approximately 215 species in the family; 
one, the Northern Pipefish, is common in the inshore marine waters of 
Massachusetts and occasionally occurs in tidal freshwater. The syngnathids 
have elongate bodies that are covered with a series of interlocking, bony 
plates, and their heads and snouts are roughly tubular, ending in a small 
mouth. Males carry the fertilized eggs and maturing larvae in a brood 
pouch until the young are well developed. The young are expelled from the 
brood pouch by the body contractions of the male. 

references. Dawson 1982 (review ofwestern North Atlantic species). 



Northern Pipefish Native 

Syngnathus fuscus Storer 1839 




identification. Northern Pipefish have elongate bodies that are cov- 
ered with interlocking bony plates. Pelvic fins are absent, and the snout is 
relatively long and tubelike. Color varies from a mottled brown to a light 
yellow-green. 

selected counts. D35-45;A2-4. 

size. Northern Pipefish rarely exceed 10 inches in length, with 6- to 
8-inch specimens being more common, especially in the upper portions of 
estuaries. 



Family and Species Accounts 229 




Northern Pipefish: coastal records are not indicated 



...... n:k« 



V 



natural history. Northern Pipefish are 
primarily an estuarine and marine species. How- 
ever, they tolerate salinities at least as low as 4 parts 
per thousand in Massachusetts and are found at the 
heads of larger coastal estuaries. These pipefishes are often found in areas 
of dense aquatic vegetation, especially in eelgrass, Zostera. Although absent 
from Massachusetts estuaries during the winter months, Northern Pipefish 
start to become common in July and peak from August to October. It has 
been shown that they migrate inshore and offshore with the season, possi- 
bly to avoid colder estuarine temperatures. 






v :- y 



distribution and abundance. Northern Pipefish are found along 
the Atlantic seaboard from Prince Edward Island, Canada, south to north- 
ern Florida. In Massachusetts, they are common in marine waters and may 
be expected to occur in the estuaries of most larger streams and rivers. 

notes. The Northern Pipefish was first described by D.H. Storer in his 
first review of Massachusetts fishes in 1839, from a specimen collected at 
Nahant. 

references. Bigelow and Schroeder 1953, Collette andHartel 1988 (gen- 
eral); Dawson 1982 (review); Fiske et al. 1968 (salinity); Lazzari and Able 
1990 (migration). 



230 Inland Fishes of Massachusetts 



Sculpin Family 

Cottidae 



The sculpins are a large family of fishes with more than 300 species in about 
70 genera. This family belongs to the so-called "mail-cheek fishes," which 
have an armorlike, bony element that extends from the cheek to the pre- 
opercle. Although sculpins are mainly marine fishes of arctic and temper- 
ate seas, one-third of the cottid species occur in freshwater. Twenty-eight 
species of the genus Cottus are widely distributed in freshwater across 
North America. Deepwater Sculpin, Myoxocephalus thompsoni, live deeper 
than any other North American freshwater fish and have been found at 
depths of 1,000 feet in Lake Superior. Sculpins are a distinctive group of 
spiny fishes with morphological adaptations reflecting their bottom-living 
habits. They have broad, somewhat flattened heads with prominent, dor- 
sally situated eyes, spines on the preopercle, and wide jaws with well- 
developed teeth. Their robust heads and bodies taper to a relatively nar- 
row caudal peduncle. Scales are generally absent, although a few may be 
found along the lateral line in some species. The fin spines are soft and flex- 
ible, and the thin pelvic spine is inconspicuous since it is embedded in soft 
tissue. Sculpins have large, fan-like pectoral fins, and the small first dorsal 
fin is separated from the second dorsal fin by a distinct notch. 

references. Jenkins and Burkhead 1993 (review); Strauss 1989 (genet- 
ics and morphology); Washington et al. 1984a, 1984b (development, 
relationships) . 



Family and Species Accounts 23 1 



Slimy Sculpin Native 

Cottuscognat us Richardson 1836 plate 54 




identification. Sculpins have a broad head, tapering body, large fan- 
like pectoral fins, a hooked preopercular spine, no scales, pelvic fins posi- 
tioned under the anterior base of the pectoral fins, and soft flexible fin 
spines. Slimy Sculpin are mottled brown to gray dorsally with saddle-shaped 
blotches that sometimes extend onto the upper sides. Breeding males are 
darker, almost black above, with a bright orange border on their first dorsal 
fin. This species is the only sculpin found in Massachusetts freshwaters. 
Another small sculpin, the Grubby, Myoxocephalus aenaeus, is common in 
local marine waters but never enters freshwater. 

selected counts. D VI-IX+14-19; A 10-14; Pel 1,3. 

size . Slimy Sculpin are relatively small fish; most adults are about 3 inches 
TL. We have seen an unusually large Slimy Sculpin, measuring 4.7 inches 
TL (96 mm SL), from a tributary to the Housatonic River in Pittsfield in 1979. 

natural history. In Massachusetts, Slimy Sculpin are known only 
from high gradient, rocky, clear, cold streams. In other parts of their range, 
they also inhabit cold-water lakes and low gradient spring-fed streams. In 
streams, Slimy Sculpin inhabit riffles and tend to stay close to the bottom, 
generally hidden in the stony substrate. Spawning takes place in the spring 
when water temperatures approach 50°F. Nests have been found in western 
Massachusetts in early May. Males establish territories around a crevice be- 
neath a log, rock, or tree root. As one or more females are enticed into the 
nest, the eggs are deposited in a cluster on the underside of the roof of the 
nest cavity and are fertilized. The eggs hatch in about four weeks. Male 
sculpins both guard the nest and care for the young. Slimy Sculpin feed pri- 



232 Inland Fishes of Massachusetts 




Slimy Sculpin: open circles indicate known localities 
where species was not found during our post- 1969 
surveys; not all solid circles were resurveyed. 



marily on bottom-dwelling invertebrates, particu- 
larly aquatic insect larvae and nymphs. Small crusta- 
ceans, fishes, and aquatic vegetation are consumed in 
lesser amounts. 



k 






*& 



distribution and abundance. In Massachusetts, Slimy Sculpin are 
common and widely distributed in suitable habitats west of the Connecti- 
cut River. East of the Connecticut River, there are small, geographically 
isolated populations in the Millers, Chicopee, and Nashua river basins. In 
1861, specimens were taken from the lower Merrimack near Lawrence, but 
this population has apparently been extirpated since they were last reported 
from the entire area in 1953. The continued presence of Slimy Sculpin in 
eastern Massachusetts depends on the protection of habitat adjoining hill- 
streams. In addition, changes in water quality, probably due to acid rain, 
have impacted a number of Slimy Sculpin populations in tributaries to the 
Quabbin Reservoir in north-central Massachusetts. 

notes. The sculpins' reputation as being detrimental to trout is un- 
founded. While sculpins are closely associated with trout in Massachu- 
setts, they have no documented negative effect on the survival of trout 
populations. In small, high-gradient streams, they are often the only fish 
species found in association with Brook Trout. Reports of Mottled Sculpin, 
Cottus bairdi, from Massachusetts may stem from the fact that Massachu- 
setts Slimy Sculpin often have four pelvic rays on one side or the other but 
are true C. cognatus in other characteristics. 



Family and Species Accounts 233 



references. Godkin et al. 1982 (identification); Halliwell 1989 (decline); 
lenkins and Burkhead 1993 (identification, review); Koster 1937 (life his- 
tory); Lyons 1990 (distribution); Mousseau et al. 1988 (reproduction); 
Schlotterbeck 1954 (Merrimack); Symons et al. 1976, Coon 1987 (tempera- 
ture); Van Vliet 1964 (ecology). 



234 Inland Fishes of Massachusetts 



Striped Bass Family 

Moronidae 



The striped bass family is a somewhat primitive group of spiny-finned 
fishes. The classification of these sea basslike fishes has changed in recent 
years, and the striped basses have been removed from the Percichthyidae 
and placed in a separate family, Moronidae. This small family consists of 
four North American species and two from the Mediterranean area of Eu- 
rope and Africa. Some are anadromous and migrate long distances. They 
have been considered to be closely related to the true sea bass family (Ser- 
ranidae), but recent research suggests that they might be more closely re- 
lated to the snooks (Centropomidae). Unlike the sea basses, members of 
this family usually have two opercular spines. The striped bass family also 
has two dorsal fins placed fairly close together and three anal spines. 

references. Johnson 1984 (relationships, development); Fritzsche and 
Johnson 1980, Waldman 1986 (development and identification). 



Key to Massachusetts Striped Basses 



la. Dorsal fins slightly joined at base by 
membrane; second anal fin spine almost 
equal in length to third; 1 opercular 
spine; no dark horizontal stripes on body. 
White Perch, Morone americana, page 
236, Plate 56. 

lb. Dorsal fins entirely separate; anal 
spines graded in length with the second 
intermediate between first and third; 2 
opercular spines; 7 or 8 dark horizontal 
stripes on body. Striped Bass, Morone 
saxatilis, page 238. 















Family and Species Accounts 235 



White Perch Native 

Morone americana (Gmelin 1789) plate 56 




identification. White Perch are relatively deep -bodied, spiny fishes 
with two dorsal fins that are just slightly connected, and three anal spines. 
Young White Perch and Striped Bass might be confused, but White Perch 
have a single opercular spine and the second anal spine is almost as long as 
the third. Adult White Perch lack the stripes found on Striped Bass. 

selected counts. D IX,I,1 1-12; A 111,9-10; Scales 8-9/46-50/10-12; 
GR 20-23. 

size . White Perch average 6 to 10 inches TL and .5 to 1 pound in weight. 
The current Massachusetts sport fish awards record is a 3-pound, 2-ounce 
fish from Wachusett Reservoir in 1994. 

natural history. White Perch tolerate a wide range of salinity and 
water temperature and live in marine, estuarine, and fresh waters. Estuarine 
populations generally spawn on sand and gravel bars in freshwater coastal 
streams during April to May. Landlocked freshwater populations spawn 
along gravel shorelines from May to June at water temperatures of 50° to 
60°F. During spawning, 10 to 50 smaller males attend a few larger females. 
Nests are not built, and the adhesive eggs are scattered over the bottom 
during one to two weeks. Parents do not take care of eggs and young. Eggs 
hatch in four to six days, depending on water temperature, and the young 
grow rapidly, reaching 1.5 to 2.5 inches TL by late summer. Their average 
life span is six to seven years, but they have been known to live for 17 years. 
White Perch tend to form schools of a few to hundreds of fish. Their prey 
vary with habitat and age. In freshwater, the young feed primarily on small 



236 Inland Fishes of Massachusetts 





^,iz,.^. 






\ 

•* 



White Perch: most of the inland populations were 
stocked. 



y>fC 

X 



j 



<•*. 



• 1— 

W 



Tj* 



plankton while the larger perch typically shift to 
crayfishes, aquatic insects, and small fishes. Studies 
indicate daily migrations into the shallows in the evening; 
they return to deeper offshore waters during the daytime. 



/;- 



SWj 



distribution and abundance. In Massachusetts, White Perch were 
originally restricted to estuarine streams and coastal freshwater ponds. 
They were introduced to many ponds statewide and are now common to 
abundant. Most inland landlocked populations are the result of stocking 
programs that started in the early 1900s. 

notes. White Perch have a strong tendency to overpopulate lakes and 
ponds, which results in large numbers of stunted individuals. When this 
happens, White Perch also compete with other species for food resources. 
In spite of its reputation as an excellent food and sport fish, White Perch are 
underused by Massachusetts anglers. 

references. Webster 1943, Alsop and Forney 1962 (food); Thoits 1958, 
Mansueti 1961, 1964a, Clayton et al. 1976, Bath and O'Connor 1982 (life his- 
tory); Holsapple and Foster 1975 (reproduction); Kellogg and Gift 1983 
(growth); Aziz 1992 (fishing, MA); Waldman 1986 (identification); Cardoza 
etal. 1993 (introductions). 



Family and Species Accounts 237 



Striped Bass . Native 

Morone saxatilis (Walbaum 1792) 



■?7T- "":'■■ : •'- , "-»' , ®W»B 





identification. Striped Bass are moderately elongate as young but can 
become quite stout as adults. Their two dorsal fins are separated by a small 
space, and they have three anal spines. Young White Perch and Striped 
Bass might be confused, but Striped Bass have two opercular spines and 
their anal spines increase gradually in length. Adult Striped Bass have an 
olive-green to bluish back that grades to silvery sides and a white belly. As 
their name implies, they have seven to eight dark horizontal stripes along 
the body. 

selected counts. D IX,I,1 1-14; A 111,11; Scales 53-65; GR 25. 

size. Striped Bass can grow to be large fish; records of over 100 pounds 
have been historically noted. Today, most specimens are from 3 to 
40 pounds. Three fishes, each weighing 73 pounds, hold the current Mas- 
sachusetts angling record. 

natural history. Striped Bass move into Massachusetts waters dur- 
ing summer months as water temperatures reach 57° to 64°F. All of the 
Striped Bass found along the Massachusetts coast and in inland rivers are 
the products of migratory populations that spawn farther south from 
the Hudson to the Chesapeake. It is unknown if Striped Bass historically 
spawned in Massachusetts rivers, and the only indication that they might 
have is the mid- 1600s description of large numbers running up the Merri- 
mack with spring Alewives, and a series of young-of-the-year stripers that 
were collected in the Parker River estuary in 1930. Stripers often migrate 
hundreds of miles inland in larger rivers. In the Connecticut and Merri- 



238 Inland Fishes of Massachusetts 




Striped Bass: open circles indicate historic records 
where the species may no longer occur; coastal 
records are not indicated. 



/>■ 



_ ~:-V "t.M 



mack rivers, one- to three-year-old fish swim into & 

New Hampshire. South of New England, stripers spawn 
in the spring, and the young stay in their native rivers or estuaries until they 
are two years old or about 1 foot long. Males and females grow at different 
rates: 15-year-old males will reach an average of 42 inches, while females of 
the same age reach 47 inches and are much heavier. The number of eggs 
produced by females also varies with age and size. A 4-pound female might 
produce 250,000 eggs although a 50-pound fish might produce somewhere 
around 5,000,000. Females do not mature until they are four to six years 
old. As might be expected, diet varies with size: young fish feed on smaller 
amphipods, shrimp, and crabs; older fishes are voracious and consume 
many larger prey, including herrings, smelt, anchovies, eels, sea-worms, 
squid, and crabs. Striped Bass may live as long as 40 years. 






distribution and abundance. Historically, Striped Bass were abun- 
dant and probably entered most of Massachusetts' larger rivers before the 
appearance of environmental changes associated with dams and pollution. 
Improvements to many of Massachusetts' fishways during the last decade 
currently allow nonreproducing stripers to migrate the length of the Con- 
necticut and Merrimack rivers into New Hampshire (Figure 3). Juvenile 
stripers are also found in the Taunton River. Striped Bass typically undergo 
natural population fluctuations that have been documented since before 
the turn of the 20th century. The changes in abundance have now been 
linked to peak years of successful reproduction followed by less successful 
years. Recently, these natural fluctuations have been compounded by 
human-induced changes that affect water quality and thus reproductive 



Family and Species Accounts 239 



and larval success. In fact, the species declined to such low levels that some 
people speculated that it might become extinct during the 1970s. Strict reg- 
ulations put in place at that time allowed the species to rebound, and by 
the late 1990s it had greatly increased in abundance. 

notes. Historical reports from the mid- 1600s note that a single man with 
a codline baited with lobster might catch from 12 to 20 bass in three hours 
in the Boston area. Likewise, two to three thousand fish could be taken in 
one tide with blocknets across a creek. Bass were considered so important 
that laws were enacted to prevent them from being used as fertilizer 20 years 
after the Pilgrims landed. In 1670, the colonists restricted all income from 
bass fished at Cape Cod to be used to establish a town school. 

references. Bigelow and Schroeder 1953, Raney 1952, Clayton etal. 
1978, Smith and Wells 1977 (abundance, life history); Frisbie 1967 (growth, 
MA); Setzler et al. 1980 (synopsis); Markle and Grant 1970 (diet); Radin 1997 
(recovery). 



240 Inland Fishes of Massachusetts 



Sunflsh and Black Bass Family 

Centrarchidae 



The centrarchid family is endemic to North America and has eight genera 
and 29 species, including such important warmwater game fishes as the 
black bass (Largemouth Bass and Smallmouth Bass), crappies, and sun- 
fishes. Due in part to their general hardiness and much-sought-after game 
fish qualities, several species have been extensively introduced and now 
have almost worldwide distributions. Within the United States, centrarchids 
have been transplanted to such an extent that their original ranges are 
sometimes in question. Centrarchids have united spiny and soft dorsal fins 
and lack opercular spines. Large opercular flaps are often present, particu- 
larly in male sunfishes, and the mouth is terminal with numerous, small, 
conical teeth. They tend to be territorial, especially during their midspring 
to midsummer breeding periods. Males build and defend nests and attract 
females with well-defined courtship displays. The nest may consist of a 
shallow pit in gravel or sand or occasionally simply a cleaned area in rubble 
or vegetation. These nests, especially in certain Lepomis species, are occa- 
sionally tightly clustered in "breeding colonies." Males and females of some 
species mate with numerous partners, and a single nest may contain the 
eggs of several females. After spawning, the males take care of the nest and 
young until several weeks after hatching. In North America, members of 
this family are some of the most economically important and sought-after 
game fishes. 

references. Avise et al. 1977, Branson and Moore 1962, Mabee 1988 
(systematics); Breder 1936 (reproductive biology); Hubbs 1955, Childers 
1967 (hybrids); Etnier 1971 (diet and hybrids); Gross 1982 (reproductive bi- 
ology); Werner and Mittelback 1981 (behavioral ecology). 



Family and Species Accounts 24 1 



Key to Massachusetts Sunfishes, Crappies, 
and Black Basses 



la. Anal fin with 3 spines (rarely 2 or 4). 
Go to 4. 




lb. Anal fin with 5 or more spines. 
Go to 2. 




2a. Dorsal fin spines 1 1 to 13; dorsal fin 
much longer than anal fin. Rock Bass, 
Ambloplites rupestris, page 246, Plate 60. 




2b. Dorsal fin spines 8 or fewer; dorsal 
and anal fins of about equal length. Crap- 
pies. Go to 3. 




242 Inland Fishes of Massachusetts 



3a. Length of dorsal fin base equal to or 
greater than distance from eye to origin 
of dorsal fin; dorsal spines 7 to 8 (some- 
times 6); body with mottled color pattern. 
Black Crappie, Pomoxis nigromaculatus, 
page 264, Plate 58. 




3b. Length of dorsal fin base less than 
distance from eye to the origin of the dor- 
sal fin; dorsal spines 5 to 6 (sometimes 7); 
body with dark vertical bars. White Crap- 
pie, Pomoxis annularis, page 262. 




4a. Fifty-eight or more scales in lateral 
series; body elongate. Black Basses. 
Go to 9. 




4b. Fifty-three or fewer scales in lateral 
series; body deep and laterally com- 
pressed. Sunfishes. Go to 5. 




Family and Species Accounts 243 



5a. Tail fin rounded. Banded Sunfish, En- 
neacanthus obesus, page 248, Plate 57. 




5b. Tail fin at least slightly forked. Go to 6. 




6a. Pectoral fins short, rounded, and 
barely reaching eye if fin is folded for- 
ward. Go to 8. 




6b. Pectoral fins long, pointed, and 
reaching beyond middle of eye if fin is 
folded forward. Go to 7. 




244 Inland Fishes of Massachusetts 



7a. Soft dorsal fin with distinct dark spot 
near base of last few rays; gill rakers on 
first arch more than twice as long as 
wide; opercular flap flexible, "squared," 
and without light-colored margin. 
Bluegill, Lepomis macrochirus, page 256, 
Plate 59. 




7b. Soft dorsal fin without large dark 
spot near base of last few rays; gill rakers 
on first arch not more than twice as long 
as wide; opercular flap stiff, rounded, and 
with red /orange spot on lower margin. 
Pumpkinseed, Lepomis gibbosus, page 
254, Plate 62. 




8a. Mouth large, maxilla often extending 
to rear margin of eye; opercular flap stiff 
and relatively short. Green Sunfish, Lep- 
omis cyanellus, page 252. 




8b. Mouth small, maxilla extending, at 
most, to midpoint of eye; opercular flap 
flexible and often long (especially in large 
males). Redbreast Sunfish, Lepomis auri- 
tus, page 250, Plate 61. 




Family and Species Accounts 245 



9a. Soft and spiny portions of dorsal fin 
connected by membrane only at base; 
dark midlateral stripe on each side; no 
scales present along the base of soft dor- 
sal fin; juveniles without tricolored tail 
(tail may be bicolored). Largemouth 
Bass, Micropterus salmoides, page 260, 
Plates 64 and 65. 




9b. Soft and spiny portions of dorsal fin 
connected well above the body; no dark 
midlateral stripe; scales present along 
base of soft dorsal fin; juveniles with tri- 
colored tail (base of fin orange, middle 
dark, outer edge white /clear). Small- 
mouth Bass, Micropterus dolomieu, page 
258, Plates 63 and 65. 




Rock Bass 

Ambloplites rupestris (Rafinesque 1817) 



Introduced 
plate 6o 




identification. Rock Bass are medium- sized centrarchids with large 
mouths that reach back to below the pupil, short rounded pectoral fins, 
10 to 12 dorsal spines and 5 to 7 anal spines. 

selected counts. D X-XII, 10-12; A V-VII,9-1 1; Scales 7/38-43/12; 
GR12. 



246 Inland Fishes of Massachusetts 




Rock Bass. 



■ . • 



UK > cV 



n 



U 



size. Individuals ranging from 6 to 8 inches TL 

are relatively common in larger bodies of water, 

but the species seldom reaches 10 inches TL. Males ...JS^- 

tend to be longer and heavier than females. 



; l/V 



r 



\ > 






natural history. Rock Bass are primarily bottom- dwelling fish. They 
inhabit portions of large streams that have cool, clear waters with extensive 
cover, such as submerged stumps or large rocks. Rock Bass are also com- 
monly found in lakes and reservoirs where the appropriate conditions of 
cool, clean water and ample cover are satisfied. In Massachusetts, spawn- 
ing probably begins in early June and continues for several weeks. The males 
build nests, often close together, in shallow water in areas of silt-free gravel 
and rocky bottoms. Females have been found to carry 2,000 to 11,000 eggs. 
Rock Bass mature at age three or four. Individuals have been found to live 
at least eight years in Massachusetts. Rock Bass feed on a variety of prey. 
The young feed mostly on zooplankton and small benthic invertebrates 
while larger individuals switch to a diet of fishes and relatively large inver- 
tebrates, such as crayfishes and dragonfly larvae. 

distribution and abundance. Rock Bass were first introduced into 
Massachusetts in 1934. Distribution in this state is primarily western, with 
the majority of records from the Hoosic and Housatonic drainages as well 
as the major tributaries to the Connecticut River. Rock Bass are locally com- 
mon in the Farmington, Deerfield, Millers, Chicopee, and Westfield rivers, 
as well as in the Quabbin and Wachusett reservoirs. 

notes. Where found, Rock Bass are an interesting game fish that put up a 
good fight on a light rod. However, the introduction of this competitive fish 



Family and Species Accounts 247 



eliminated a population of the Roanoke Bass, Ambloplites cavifrons, and a 
Brook Trout fishery in Virginia. 

references. Cashner and Jenkins 1982 (systematics); Gross and Nowell 
1980 (reproductive biology); Keast 1977b (behavioral ecology); Keast and 
Webb 1966 (diet); Noltie 1988 (growth); Trautman 1981 (general); Woronecki 
1966 (life history, Quabbin Reservoir); Jenkins and Burkhead 1993 (VA). 



Banded Sunfish 

Enneacanthus obesus (Girard 1854) 



Native 
plate 57 




identification. Banded Sunfish are small (usually less than 3 inches 
TL), stout-bodied fishes, and they are the only local sunfish with rounded 
tail fins and short-round pectoral fins. They are olive- green to brown with 
numerous small bronze, silver- green, and light blue spots on the body, and 
five to eight dark vertical bars. 

selected counts. D VIII-IX, 11-12; A 111,10-11; Scales 5/30-33/10-11; 
GR 13-14. 

size. Banded Sunfish rarely exceed 2 to 3 inches TL. The largest Massachu- 
setts specimen that we have seen measures 4.25 inches TL (80 mm SL). 

natural history. Banded Sunfish live in quiet backwaters, swamps, 
and ponds. They are frequently associated with heavy aquatic vegetation 
and have been observed to thrive in naturally acidic waters. They appear to 
have a protracted breeding season, beginning in late spring and extending 



248 Inland Fishes of Massachusetts 







Banded Sunfish: open circles indicate known locali- 
ties where species was not found during our post- 
1969 surveys; not all solid circles were resurveyed. 



■,fi£- 



-•5 \9 






/±J-~-"^!/ 



k 



though midsummer. During this period, Banded ; ^ A. 

Sunfish aggressively defend small territories. Spawn- </ 

ing occurs in small nests constructed at the base of aquatic vegetation 
or occasionally well above the bottom in the vegetation itself. Banded Sun- 
fish usually live three to four years, although five-year-old specimens have 
been recorded. They feed on a wide variety of small aquatic invertebrates, 
including cladocerans, copepods, dipterans, and amphipods. The relative 
size and seasonal abundance of prey greatly influence their diet. In the lab- 
oratory, they readily feed on small fishes and crayfishes. 



distribution and abundance. Banded Sunfish inhabit the Atlantic 
coast from New Hampshire to Alabama. The species is widespread in most 
of eastern Massachusetts but known only from a few sites on Cape Cod. We 
found it at two locations on Martha's Vineyard in 1988, but not on Nan- 
tucket. In the inland portion of the state, Banded Sunfish have been seen 
only in the upper Chicopee Drainage (Burnshirt and Ware rivers), Nashua 
River, and the upper Millers River drainages. These Chicopee and Millers 
populations are most likely the result of stream capture with the Merrimack 
Basin. This species is still common in the proper habitat but has declined 
with urban sprawl when small, swampy wetlands were drained. 

notes. The Banded Sunfish was first described from specimens collected 
at Hingham and Holliston, Massachusetts, by C. Girard in 1854. 

references. Breder and Redmond 1929 (reproduction); Graham and 
Felly 1985 (hybrids); Gonzales and Dunson 1989 (pH tolerance); Graham 



Family and Species Accounts 249 



and Hastings 1984 (habitat); Harrington 1956 (reproduction); Lima 1986 
(ecology); Sweeny 1972 (systematics). 



Redbreast Sunfish Native 

Lepomis auritus (Linnaeus 1758) plate 6i 



identification. Redbreast Sunfish have united dorsal fins, three anal 
fin spines, short and rounded pectoral fins (not reaching to the front of the 
eye when folded forward) (see key Figure 6a), less than 13 short thick gill 
rakers, and a relatively large mouth that reaches well past the anterior edge 
of the eye. Adults (particularly males) have elongate, dark-blue or black op- 
ercular flaps that lack a red or orange posterior margin. Redbreast Sunfish 
are much more elongate and less deep-bodied than most other local sun- 
fishes. The body is olive-green to yellow-tan, the belly is red to yellow, and 
several light blue, irregular horizontal stripes cross the cheeks and gill cov- 
ers. The fins, particularly the tail and pelvic fins, are often red or orange. 

selected counts. D X,10-12; A 111,9-10; Scales 7/42-46/14; GR 10-13. 

size . Individuals 4 to 8 inches TL have been commonly recorded. The 
largest Massachusetts specimen that we have measured is about 6 inches 
TL(126mmSL). 

natural history. Redbreast Sunfish generally live in ponds, lakes, 
or the slow-moving sections of streams and rivers. They are often found in 
clean water with rocky bottoms and seem to avoid heavily vegetated areas. 
The spawning season is from late spring through midsummer. In rivers and 
streams, Redbreast Sunfish usually make nests in sheltered areas, near 



250 Inland Fishes of Massachusetts 




Redbreast Sunfish: open circles indicate known lo- 
calities where species was not found during our post- 
1969 surveys; not all solid circles were resurveyed. 



A"S>' 



)> V 



,/,■ 



......?Z\*»J~f 



underwater objects such as rocks and fallen trees. 

In ponds and lakes, the males dig nests in areas of gravel 

and sand bottoms. These nests are often tightly grouped. 

Medium- sized females have been recorded to contain 500 to 6,500 eggs. 

Diet consists of a wide variety of larval and adult aquatic insects, including 

mayflies, caddisflies, midges, flies, mosquitoes, beetles, and dragonflies. 

Scuds, aquatic sowbugs, mollusks, and small fishes are occasionally eaten. 



J\ 



distribution and abundance. In Massachusetts, Redbreast Sunfish 
have a scattered distribution. D.H. Storer (1839) mentioned that this spe- 
cies was common, but currently we have found it to be relatively common 
only in a few locations: the upper Charles and Sudbury rivers, the Quabbin 
Reservoir, and the main stems of the Connecticut and Merrimack rivers. In 
other Massachusetts drainages, this species is not common, and it is rare or 
absent in the southeastern portion of the state and most hillstream areas. 
It is probable that this species has declined since the mid- 1800s due to 
changes in water quality or behavioral interactions with introduced spe- 
cies, especially the Bluegill. 

notes. The Longear Sunfish, Lepomis megalotis, has often been reported 
from Massachusetts, but we have never seen a specimen. These reports are 
probably the result of misidentifications of adult male Redbreast Sunfish 
or incorrect use of the common name. In early references, Redbreast Sun- 
fishes were occasionally referred to as Longear Sunfish (Kendall 1908). The 
Redbreast Sunfish is known to hybridize with Pumpkinseeds, Green Sun- 
fish, Bluegills, and Warmouths. 



Family and Species Accounts 251 



references. Bass and Hitt 1975 (ecology); Breder and Nigrelli 1935 (so- 
cial behavior); Davis 1972 (reproduction). 



Green Sunfish introduced 

Lepom is cyanellus Rafinesque 1819 



identification. Green Sunfish have united dorsal fins, three anal 
fin spines, short and rounded pectoral fins not reaching past the posterior 
margin of the eye if folded forward (see key Figure 6a), and a large mouth 
that reaches well past the anterior margin of the eye or even past the poste- 
rior margin in large adults. The opercular flap is dark with a distinct light- 
colored margin, and is rigid all the way to its posterior edge. The dorsal, 
anal, and caudal fins have a yellow- cream margin, and there are dark 
blotches at the base of the soft dorsal fin and the anal fin. A series of blue- 
green spots and wavy lines spread from the eye to the gill cover. 

selected counts. D X,10-12; A III, 9-10; Scales 7/45-53/15; GR 19-20. 

size. Green Sunfish are generally small- to medium-sized fishes that typi- 
cally reach 4 to 6 inches TL. 

natural history. In other areas, Green Sunfish tolerate a wide range 
of water conditions ranging from cool, clear hill streams and brooks to 
warm, turbid lowland backwaters. They appear to do best, however, in 
small, slow-moving streams with large amounts of aquatic vegetation or 
other cover. In Massachusetts, the breeding season is probably from late 
spring to midsummer. As in other sunfishes, the males build small nests 
in shallow water, often closely grouped. Green Sunfish feed on a variety of 
aquatic invertebrates. Large benthic prey, such as larval dragonflies, usually 



252 Inland Fishes of Massachusetts 



:'t j 




Green Sunfish. 



) 



comprise a large portion of the diet. Fishes and 
snails are also frequently eaten. 



distribution and abundance. The Green Sunfish 
is not native to Massachusetts. The exact date of its first introduction into 
Massachusetts is unknown, but the species may have been accidentally in- 
troduced with shipments of Bluegills. Green Sunfish have a limited range in 
this state. Previously, they were common only in Nagog Pond, Acton, but 
1998 MDFW surveys by T. Richards show that the species is common at 
several sites in the Ipswich Drainage. Photographic evidence of a single 
specimen documents this species in the Buzzards Bay Drainage. Unverified 
reports suggest that this species may be also found in the Taunton, Black- 
stone, and Quinebaug drainages. 

notes. Green Sunfishes are known to hybridize with Bluegills and Pump- 
kinseeds, and with Longear, Lepomis megalotis, Redear, Lepomis microlo- 
phus, and Orangespotted sunfishes, Lepomis humilis. 

references. Hunter and Hasler 1965 (reproduction and associations); 
McKechnie and Tharratt 1966 (review); Sadzikowski and Wallace 1976 
(diet). 






Family and Species Accounts 253 



Pumpkinseed Native 

Lepomis gibbosus (Linnaeus 1 758) plate 62 



identification. Pumpkinseeds have three anal fin spines, a slightly 
forked tail, united dorsal fins, and fewer than 45 lateral-line scales. The long 
and pointed pectoral fins (when folded forward they reach the front of the 
eye) (see key Figure 6b) distinguish Pumpkinseeds from all other local sun- 
fishes except Bluegills. Pumpkinseeds have a prominent orange-red spot 
and light margin on the rounded and inflexible opercular flap and lack a 
dark spot on the soft dorsal fin. They have fewer than 12 gill rakers that are 
relatively short, stout, and often crooked. However, the gill rakers of juve- 
nile Pumpkinseeds and Bluegills appear similar when the fishes are less 
than 1 inch TL. The body often has 6 to 10 darker vertical bands along the 
body and many pale yellow, orange, golden, and olive spots on the sides of 
the body and head. A series of wavy, horizontal, blue bars extends from the 
side of the head onto the lower sides of the body. Male coloration becomes 
intense during breeding. 

selected counts. D X,l 1-12; A 111,9-10; Scales 6/38-44/13; GR 1 1-15. 

natural history. Pumpkinseeds are generally associated with areas of 
aquatic vegetation in lakes, ponds, marshes, and slow-moving portions of 
streams. Spawning is temperature dependent and usually begins in late 
spring and often lasts through midsummer. Males dig small depressions 
in shallow water, usually in open areas with sand or gravel bottoms, and 
defend territories that are often only a few times the size of the actual nest. 
Nests have been known to contain 1,500 to 14,000 fry, and females between 



254 Inland Fishes of Massachusetts 










'-•? 



Iw 



Pumpkinseed. f • r" / *L ^ 4 %** i tj\ /-^ .. . 

4 and 5 inches TL have been known to carry 600 

to 3,000 eggs. This species may live up to eight 

years. Its diet consists of a wide range of aquatic in- ; Jv,* 

vertebrates, especially those dwelling on the bottom or in 

vegetation. In some populations, large juvenile and adult Pumpkinseeds 

feed almost exclusively on snails by using their molar-like pharyngeal teeth. 






size . Individuals 4 to 5 inches TL are common, and occasionally specimens 
up to 10 inches TL are encountered. 

distribution and abundance. In Massachusetts, Pumpkinseeds are 
common and live in virtually all parts of the state where quiet, vegetation- 
filled waters are present. They are found on Nantucket but not on Martha's 
Vineyard. 

notes. Pumpkinseeds and Bluegills are two of the most common fishes 
in Massachusetts and are probably the first species caught by most begin- 
ning anglers. Pumpkinseeds are known to hybridize with Bluegills and 
Warmouths, Lepomis galosus, and with Green, Redbreast, Orangespotted, 
Lepomis humilis, and Longear sunfishes. 

references. Colgan and Gross 1977 (behavior); Hanson and Qadri 1984, 
Keast 1978, Mittelbach 1984 (diet); Miller 1963 (behavior). 



Family and Species Accounts 255 



Bluegill 

Lepomis macrochirus (Rafinesque 1819) 



Introduced 
plate 59 




identification. Bluegills have three anal fin spines, a slightly forked 
tail, united dorsal fins, and fewer than 46 lateral-line scales. The long and 
pointed pectoral fins (when folded forward they reach the front of the eye 
(see key Figure 6b) distinguish Bluegills from all other local sunfishes ex- 
cept Pumpkinseeds. Bluegills have a prominent dark spot on the soft dorsal 
fin and a relatively long, dark, and flexible opercular flap. The opercular flap 
is dark blue to black to its margin. Bluegills have more than 17 relatively 
long, thin, and straight gill rakers. Overall color ranges from olive-green to a 
dusky, yellow-brown and bronze, with breeding males becoming slate blue 
on the head. The breast ranges from a light yellow- cream to a bright orange- 
red, to mahogany in breeding males. 

selected counts. D X,10-12; A 111,10-12; Scales 7/39-46/14; GR 18-20. 

size . Individuals 5 to 7 inches TL are frequently encountered, and large 
specimens up to 12 inches TL are not uncommon in some populations. A 
large Bluegill, weighing 2 lbs. 1 oz., caught at South Athol Pond in 1982, is 
the current angling record for a sunfish in Massachusetts. 

natural history. Bluegills typically live in warm, slow-moving portions 
of rivers and streams, ponds, lakes, and marshes. They have been extensively 
transplanted and may be found in almost every aquatic habitat in the state. 
The young usually spend much of the time inshore, often in and near aquatic 
vegetation. The adults spend much of the time in open water, well up in the 



256 Inland Fishes of Massachusetts 




.. .. _ ;f"-xjjm ■ \ f~ I ...» - i+itf/- : , / 



\i i-Zj.J?.A A.zA.i.Z...«s£.*.i.*C -" - ■'•■•■•■•■ 






rfif> 



\vkV .-. i "' v " 



l«U*! 



Bluegill: open circles indicate known localities 
where species was not found during our post- 1969 
surveys; not all solid circles were resurveyed. 



water column or loosely associated with large ob- & - 

jects such as fallen trees, docks, and areas of aquatic 
vegetation. Bluegills start spawning in Massachusetts in 
mid-May and continue through midsummer, usually having several peak 
breeding periods during this season. The males build nests in shallow water 
or simply take over nests constructed by other species. These nests are often 
grouped closely together, forming loose breeding colonies. Larger females 
may carry over 40,000 eggs. Bluegills normally live about 5 years, but there 
are records of 11 -year-old fishes. The young feed primarily on small ben- 
thic invertebrates and zooplankton. Adults feed on a wide variety of food 
sources, including zooplankton, small fishes, benthic invertebrates, and 
terrestrial insects from the water's surface. Aquatic vegetation is commonly 
ingested but is probably eaten only by accident when Bluegills are feeding 
on prey associated with the vegetation. 






distribution and abundance. Bluegills are not native to Massachu- 
setts. They were first introduced in 1917 and are now found in virtually all 
areas of the state, including Martha's Vineyard. The Bluegill is one of the 
most abundant fishes in the state's warmwater habitats. 

notes. Bluegills have been known to hybridize with Green and Redbreast 
sunfishes and with Largemouth Bass. In many Massachusetts localities, 
Bluegills have rapidly overpopulated the area, with the result that most in- 
dividuals are stunted. 



Family and Species Accounts 257 



references. Avila 1976 (reproductive biology); Dominey 1981 (reproduc- 
tive ecology); Gerking 1962 (trophic biology); Keast 1977b (diet); Mayhew 
1956 (reproduction); Maciolek 1984 (hybrids). 



Smallmouth Bass introduced 

Micropterus dolomieu Lacepede 1802 plates 63, 65 




identification. Smallmouth Bass are similar to Largemouth Bass. In 
Smallmouth Bass, the two dorsal fins are only partially separated by a notch 
and numerous small scales extend onto the base of the anal and dorsal fins. 
The posterior end of the upper jaw of the Smallmouth Bass reaches only 
mideye, even in the largest individuals. Adults have a series of 8 to 12 dark, 
irregular, vertical bars along the sides of the body, and three dark stripes 
radiate from the eye across the cheeks and gill covers. The tails of young 
Smallmouth Bass are distinctly tricolored, and they lack a prominent mid- 
lateral body stripe (see Plate 65). 

selected counts. D X,13-15; A III, 10-12; Scales 13/70-78/20; GR 8-11. 

size . Smallmouth Bass generally range from 8 to 13 inches, with larger 
individuals less common. The current Massachusetts state record is 
8.1 pounds and was taken from Wachusett Reservoir in 1991. 

natural history. Smallmouth Bass prefer clear, cool lakes and larger 
streams that have abundant shelter in the form of rocky areas. They gener- 
ally live in cooler and cleaner waters than the Largemouth Bass. Spawning 
occurs in late spring and early summer. Males construct large, rock-lined 
nests, up to 2 to 3 feet in diameter, often near a large underwater object 
such as a boulder or fallen tree. The male courts and spawns with several 



258 Inland Fishes of Massachusetts 



<^rL^\ 




Smallmouth Bass. 

females. Each female may carry from 5,000 to 
14,000 eggs. Smallmouth Bass mature at 3 to 
4 years and may live to 15 years. The diet of the .... J.%<* .IS 

Smallmouth Bass is variable, depending heavily on prey 
availability. In general, small individuals feed on a variety of aquatic inverte- 
brates, primarily zooplankton, and occasionally small fish. As they get larger, 
Smallmouth Bass have a diet that includes more crayfishes and fishes, and, 
in some populations, Smallmouth Bass eat crayfishes almost exclusively. 



U. :i.y 



distribution and abundance. Smallmouth Bass are not native to 
Massachusetts. The closest parts of their native range to Massachusetts are 
in areas west of the Hudson Drainage. The range of this species has been 
greatly extended through its introduction as a game fish. In Massachusetts, 
the Smallmouth Bass was first reported in 1850. Since the middle of this 
century, these bass were stocked in many of the state's reservoirs, lakes, 
and streams and can be considered locally common. In general, they are 
more common in the western and southeastern portions of the state. 

notes. Smallmouth Bass are one of the most sought-after game fishes in 
the United States and are popular in Massachusetts. They are known to hy- 
bridize with Spotted Bass, Micropterus punctulatus, and rarely with Large- 
mouth Bass. 

references. Hubbs and Bailey 1938, Jenkins and Burkhead 1993 (gen- 
eral); MacLean et al. 1981 (age-class and temperature); Mirick 1988 (MA 
introductions, fishing). 



Family and Species Accounts 259 



Largemouth Bass introduced 

Micropterus salmoides (Lacepede 1802) plates 64, 65 




identification. Largemouth Bass are similar to Smallmouth Bass. In 
the Largemouth Bass, the notch between the hard and soft portions fully 
separates the two dorsal fins, while in the Smallmouth Bass, the two fins are 
clearly joined at the base. Largemouths lack small scales along the base of 
the soft dorsal and anal fins, and in large fish the upper jaw of this species 
extends well past the eye. The relative length of the jaw, however, changes 
as the individual grows, and it is difficult to use as an identification charac- 
ter in small specimens. Adults and juveniles have a dark horizontal midlat- 
eral stripe, and the tails of young are bicolored, not tricolored as in the 
Smallmouth Bass (see Plate 65). 

selected counts. D X-XI,12-13; A 111,10-12; Scales 7/60-68/15; 
GR8-10. 

size. Adult Largemouth Bass commonly range from 10 to 16 inches TL 
with larger individuals being fairly common. The current Massachusetts 
state record, caught at Sampson Pond in 1975, weighed 15 lbs. 8 oz. 

natural history. Largemouth Bass are typically found in quiet, warm- 
water areas and are often associated with floating and submerged vegeta- 
tion. Spawning, which lasts from midspring to early summer, usually starts 
when water temperatures reaches 60°F. Males dig large nests, often 2 to 
3 feet in diameter, in areas with sand or gravel bottoms. These nests and 
surrounding territories are defended by the males. Multiple spawnings in a 
given nest are common, because each male will court several females over 
the course of the reproductive season. Likewise, females will often spawn 
with more than one male at more than one site. Hatching occurs in less 



260 Inland Fishes of Massachusetts 




• 



Largemouth Bass. 



than a week, usually in two to five days. The fry w J I • • 

form large schools and are casually guarded by the 

male. The number of fry per nest can range from 700 . ../'Jw & 

to 11,500, with most nests containing 5,000 to 7,000 young. 

Growth can be rapid, and the young-of-the-year often reach 6 inches or 

more by the end of their first growing season. The young feed primarily on 

small aquatic invertebrates but will readily take small fishes if available. As 

Largemouth Bass increase in size, fish become a greater part of their diet. 

However, they eat, or try to eat, almost anything that swims by, including 

frogs, small mammals, and birds. Largemouth Bass have been known to 

live at least 15 years. 



V 



/' 

" 1 



distribution and abundance. Largemouth Bass are native to a large 
area of central and southeastern North America. Largemouth Bass were 
first introduced into Massachusetts prior to 1862. Due to extensive public 
and private stocking programs, Largemouth Bass can be found in almost 
any body of water in the state. It is one of the most common species in 
many of Massachusetts' warmwater habitats. 

notes. The Largemouth Bass ranks high on the list of favorite warmwater 
game fish in Massachusetts. It is a voracious feeder that readily takes both 
natural and artificial baits. It attains a large size and is an excellent sport 
fish. Largemouth Bass have been documented to hybridize rarely with 
other members of the genus Micropterus and Bluegills. 

references. Emig 1966 (general biology); Hoyle and Keast 1987 (feeding 
behavior); Hubbs and Bailey 1940 (general summary); Maciolek 1984 (hy- 
brids); McCaig et al. 1960 (Quabbin). 



Family and Species Accounts 261 



White Crappie 

Pomoxis annularis (Rafinesque 1818) 



Introduced 




identification. White Crappies have compressed, deep bodies, large 
mouths, sloping concave foreheads, usually five or six dorsal spines, and six 
or seven anal fin spines. The length of the base of the dorsal fin is usually 
less than three-quarters of the length from the eye and the origin of the dor- 
sal fin (see key Figure 3b). White Crappies have a regular series of diffuse 
vertical bars along the sides of the body. 

selected counts. D V-VI(VII), 14-15; A VI-VII, 17-18; Scales 6/34-44/ 
13-14; GR 28-32. 

size . Individuals 8 to 13 inches in total length are common in many 
populations. 

natural history. White Crappies are found in lakes, ponds, marshes, 
and slow-moving sections of rivers and streams. They are more tolerant of 
siltation and turbid water than Black Crappies and are less dependent on 
cover. Spawning occurs from midspring to early summer. Females carry 
from 1,000 to 200,000 eggs, depending on their size. Young White Crappies 
feed primarily on zooplankton, which they catch in midwater. Larger indi- 
viduals also feed on midwater organisms, including relatively small fishes 
and aquatic invertebrates. White Crappies are most active during the twi- 
light hours and at night. 

distribution and abundance . White Crappies are native to the Mis- 
sissippi River Basin, the Great Lakes region, and the western Gulf Coast 



262 Inland Fishes of Massachusetts 







White Crappie. 



*--\. 






■■Hy 

> V / \ ! _. ..>, ) ■.) \J / \' ' •' 

drainages. In Massachusetts, the White Crappie's ( ) i/J 

known range is limited to the Connecticut River in 
the vicinity of the Easthampton Oxbow. They were first ^,j.%j^j1j 
introduced to this state in 1930. There is also an undocu- 
mented report from Lake Chauncy, Westborough, but we have not seen 
specimens from that area. The White Crappie has always been an uncom- 
mon species in Massachusetts, but it appears to have continuously repro- 
duced in this small area of the state since its introduction. 



> 



'••■..'■ 



,. .... </ 



% 



notes . The White Crappie is known to hybridize with the Black Crappie. 

references. Goodson 1966 (general biology); O'Brien et al. 1986, Wright 
and O'Brien 1984 (feeding), Mugford 1969 (Lake Chauncy). 



Family and Species Accounts 263 



Black Crappie introduced 

Pomoxis nigromaculatus (Lesueur 1829) plate 58 



identification. Black Crappies have compressed deep bodies, large 
mouths, sloping concave foreheads, usually seven or eight dorsal spines, 
and six or seven anal fin spines. The length of the base of the dorsal fin is 
approximately equal to the distance between the eye and the origin of the 
dorsal fin (see key Figure 3a). Black Crappies have a color pattern that is 
characterized by irregular mottling of dark spots and patches, while White 
Crappies have regular series of diffuse vertical bars along the sides of the 
body. During the breeding season, males often become much darker, al- 
most black in color. 

selected counts. D (VI) VH-VIII, 15-18; A VI-VII, 16-18; Scales 7/36- 
42/12; GR 27-29. 

size. Black Crappies are often encountered in the 8- to 12-inch TL range 
with larger individuals not uncommon. The current Massachusetts state 
record weighed 4 lbs. 10 oz. and was caught in Jakes Pond, Plymouth, in 
1980. 

natural history. Black Crappies are found in rivers and streams, usu- 
ally in quiet backwaters or deeper areas, and in ponds and lakes. They are 
often associated with cover, such as overhanging trees, submerged brush, 
docks, and aquatic vegetation. Black Crappies often form schools, but 
larger individuals are somewhat solitary. Spawning occurs from midspring 
to early summer when water temperatures are greater than 68°F. Shallow 
nests, 6 to 8 inches in diameter, are constructed by the males in areas with 



264 Inland Fishes of Massachusetts 








LfV *:,/f?|,-1^ Vi L Ji ^V5\VV s' ■■■■->■ 

\\M <^Al .t. < *- t ! > ^iv^y ♦-.• 



Black Crappie. 




"f> 






sandy substrates. The nests, which may be clus- 
tered, are usually in water that is 1 to 9 feet in 
depth. Females have been found to carry from 1 1,000 
to 180,000 eggs. During the spawning season, a female 
will spawn several times, laying eggs in the nests of a number of different 
males. Black Crappies forage primarily in midwater; large individuals feed 
almost entirely on other fishes. Small Black Crappies typically feed on 
aquatic invertebrates but will readily eat young fishes. Populations of 
stunted Black Crappies are common in locations lacking sufficient numbers 
of forage fishes. Black Crappies feed extensively during the twilight hours 
and at night, when they leave areas of cover and move into open water. 

distribution and abundance. Black Crappies are native to a large 
area of the United States and southern Canada outside of New England. In 
Massachusetts, Black Crappies were first introduced in 1910 and were ex- 
tensively stocked until 1940. They are common in many portions of the 
state, particularly in the central and eastern areas. 

notes. Black Crappies, also known as "Calico Bass," are a popular game 
fish in many parts of the United States. The larger Black Crappies that are 
angled in Massachusetts typically come from large waterbodies, such as the 
Quabbin Reservoir and Onota and Quinsigamond lakes. Black Crappies hy- 
bridize with White Crappies. 

references. Goodson 1966 (general biology); Hanson and Qadri 1984 
(diet); Keast 1968 (trophic biology) . 



.S\ 



Family and Species Accounts 265 



Perch Family 

Percidae 



The perch family is found in the temperate freshwaters of the Northern 
Hemisphere. These spiny-rayed fishes are placed in nine genera with about 
165 species. The American members can be divided into three groups 
(tribes): the perch group, Percini, which contains the Yellow Perch; the tribe 
Luciopercini, which is represented locally by the introduced Walleye; and 
the darters, Etheostomatini, which are found only in North America. There 
are over 145 darter species, with some species still undescribed. Many dart- 
ers inhabit limited geographic areas and are listed as threatened or endan- 
gered. The controversy over the possible extinction of the Snail Darter, Per- 
cina tanasi, during construction of the Tellico Dam in Tennessee brought 
this group of fishes to the attention of the world. Most darters are colorful. 
In contrast, the two species found in Massachusetts are somewhat drab 
and do not exhibit the yellows, reds, greens, and blues so common in many 
other darter species. 

references. Collette et al. 1977 (biology); Collette and Banarescu 1977 
(systematics, zoogeography); Kuehne and Barbour 1983, Page 1983 (re- 
views), Page 1985 (reproduction); Hokanson 1977 (temperature require- 
ments). 



Key to Massachusetts 
Perches and Darters 



la. Mouth small, maxilla reaching only 
front margin of eye. Darters. Go to 3. 




lb. Mouth large, maxilla extending to 
midpoint of eye or beyond. Go to 2. 



266 Inland Fishes of Massachusetts 




2a. Anal fin with 6 to 8 soft rays; teeth 
in lower jaw all of approximately equal 
height. Yellow Perch, Perca flavescens, 
page 272, Plate 66. 



2b. Anal fin with 12 to 13 soft rays; typi- 
cally two canine teeth, much larger than 
surrounding teeth, at anterior tip of 
lower jaw. Walleye, Stizostedion vitreum, 
page 274. 




3a. Complete lateral line along midline 
of body; anal fin with 1 thin spine. Tessel- 
lated Darter, Etheostoma olmstedi, page 
270, Plate 68. 




3b. Incomplete lateral line placed high 
on body, end about midbody; anal fin 
with 2 thin spines. Swamp Darter, 
Etheostoma fusiforme, page 268, Plate 67. 




Family and Species Accounts 267 



Swamp Darter Native 

Etheostoma fusiforme (Girard 1 854) plate 67 



identification. Swamp Darters have an incomplete lateral line that 
curves upward within three scales of the first dorsal fin and two anal spines. 
The body is brownish tan with about 10 dark-brown blotches that often 
merge into a continuous band along the midside and a small spot at the 
base of the tail. Overall coloration varies; fishes from weedy and tannic 
waters are quite dark, while those from sandy-bottomed clear ponds are 
somewhat lighter in color. 

selected counts. D IX-XI,8-13; A 11,5-10; Scales 3/51-54/7-10; GR 7-8. 

size . Adults range from 1 to 2 inches TL. The largest Massachusetts speci- 
men that we have measured is 47 mm SL. 

natural history. Swamp Darters typically inhabit still or slow-flowing 
water where vegetation is abundant over mud and detritus bottoms. On 
Cape Cod and the Islands, however, they are found in clear-water ponds 
with only moderate vegetation. In Massachusetts, Swamp Darters probably 
spawn between mid-April and mid-May. Aquarium observations show that 
male darters court females by mounting them and making contact with 
their pelvic fins. If the female accepts a male, she leads him into aquatic 
vegetation where eggs are laid, one at a time, and fertilized on the plants. 
Eggs hatch in 8 to 10 days, and the young reach adult size by fall. It is be- 
lieved that most Swamp Darters breed only once and die before their sec- 
ond breeding season. Swamp Darters often rest on aquatic vegetation and 
dart out to capture small aquatic organisms. Copepods seem to be their 
most common prey. 



268 Inland Fishes of Massachusetts 




Swamp Darter. 



distribution and abundance. In Massa- 
chusetts, Swamp Darters are found in all major 
drainages in the eastern part of the state, on Cape .,.J^»j* 

Cod, Nantucket, and Martha's Vineyard. Swamp Darters 
are still common in many areas of eastern Massachusetts. Their overall 
distribution, however, has been reduced due to development of the large 
eastern cities and towns. The Nantucket population in Gibbs Pond was pre- 
sumed extirpated in 1935, but it still persists today, with small numbers of 
specimens being found in 1956, 1981, 1987, and 1995. The Martha's Vine- 
yard populations, although never reported previously, are common in 
Seths and Old House ponds. 



notes . The Swamp Darter was originally described from tributaries of the 
Charles River near Framingham, Massachusetts, by C. Girard in 1854. The 
Cape Cod and Nantucket populations were thought to be distinct and were 
named as subspecies called metae-gadi (Cape Cod) and insulae (island) by 
C. Hubbs and M. Cannon in 1935. However, studies by B.B. Collette show 
that these populations do not deserve subspecific rank. 

references. Collette 1962 (systematics, distribution, life history); Hubbs 
and Cannon 1935 (systematics and description of Cape Cod and Nantucket 
populations); Schmidt and Whitworth 1979 (distribution, New England). 



Family and Species Accounts 269 



Tessellated Darter Native 

Etheostoma olmstedi Storer 1842 plate 68 



identification. Tessellated Darters have a continuous lateral line along 
the midbody that does not arch upward, and they have a single anal spine. 
Nonbreeding Tessellated Darters are colored sandy-tan with several dark 
saddle-like marks and 9 to 10 dusky, lateral spots that often suggest the let- 
ters x, y, or z. Males usually have a dark blotch on the membrane between 
the first and second dorsal spines, become quite black, and develop swollen 
white tips on the pelvic fins when breeding. 

selected counts. D XIII-XVI, 12-15; A 1,7-8; Scales 4-6/42-48/6-9; 
GR 7-9. 

size . Adults are usually 2 to 3 inches TL. Raney and Lachner (1943) re- 
ported an 88 mm SL specimen from Massachusetts, and we have collected 
a few specimens over 4 inches TL (83 mm SL). 

natural history. Tessellated Darters prefer moving water and, unlike 
Swamp Darters, are seldom found in lakes or ponds. This darter frequents 
areas with rubble, sand, or mud bottoms that usually have some vegetation. 
They often sit motionless, propped up on their pelvic fins, on the bottom 
or on rocks and then make abrupt, quick darts when feeding or disturbed. 
Underwater objects, usually rocks or logs, are required for spawning. In 
early spring, a male selects and cleans a nest site on the underside of an 
object and then attracts a female. He leads the female to the nest site, and 
they spawn while hanging upside down under the object. A male will spawn 
several times with one or more females, until a large cluster of eggs is de- 
posited. After egg-laying, males are solely responsible for the eggs, which 



270 Inland Fishes of Massachusetts 



<:-l -j 




*£**& <*v 



Vf; W\( 




\i., 



Tessellated Darter: open circles indicate known lo- 
calities where species was not found during our post- 
1969 surveys; not all solid circles were resurveyed. 

they vigorously defend. The male also spends con- 
siderable amounts of time cleaning the eggs, either by 
fanning them with his tail or moving over them with his 
pelvic fins. After the eggs hatch, the young remain in the spawning area 
until they are about 1.5 inches TL. Males tend to grow faster and larger than 
females, and this size difference relates to their ability to defend the nest 
site. This species is short-lived, and most individuals die after their third 
summer, although a few survive into their fourth winter. Tessellated Darters 
feed mainly on the larvae of midges and other flies; however, they may 
switch to other food, such as caddisflies, later in the season. 



/ 



distribution and abundance. In Massachusetts, this darter is 
common in most of the Connecticut and Blackstone river basins, in the 
southeastern parts of the state, and on Martha's Vineyard. It is rare in 
the northeast drainages, where only a few specimens have been found 
in the Merrimack River Drainage. This darter is absent from the Hoosic, 
upper Deerfield, Charles, and Nantucket drainages. 

notes. In Massachusetts, this species has often been mistakenly called the 
"Johnny Darter." The true Johnny Darter, E. nigrum, is widespread outside 
of New England. The Tessellated Darter was originally described in 1842 
from the Connecticut River near Hartford by D.H. Storer, an early Massa- 
chusetts ichthyologist. 

references. Atz 1940 (reproduction); Chapleau and Pageau 1985 (sys- 
tematics); Cole 1967 (systematics and distribution); Constantz 1985 (behav- 



Family and Species Accounts 271 



ior); Raney and Lachner 1943 (age and growth); Layzer and Reed 1978 
(food, age, growth); Tsai 1972 (life history). 



Yellow Perch 

Perca flavescens (Mitchill 1814) 



Native 
plate 66 




identification. Yellow Perch have a short anal fin (six to eight rays) and 
lack canine teeth in the jaws. The first dorsal fin membrane has a dusky 
blotch between the last four spines, and the pelvic and anal fins of mature 
adults are brilliant red-orange. Small young are more silvery, while juve- 
niles are overall green, without red on lower fins. 

selected counts. D XII-XV+1, 13-15; A 11,7-8; Scales 6-8/53-64/13-14; 
GR 19-23. 

size . Most Yellow Perch taken by local anglers are 8 to 12 inches TL, and 
any specimen in excess of 12 inches and weighing over 1 pound would be 
considered large. The Massachusetts state record is a 2-pound, 12-ounce 
Yellow Perch from South Watuppa Pond in 1979. 

natural history. Schools of Yellow Perch commonly occur in both 
clear and weedy areas of lakes and ponds and in the slow-moving parts of 
larger streams and rivers. Young inhabit weedy shallows, while adults pre- 
fer rock ledges or submerged bars in deeper waters. Spawning occurs at 
night in shallow weedy areas during early April and May, when water tem- 
peratures are between 44° and 54°R Gravid females are attended by as many 
as 15 to 25 males. Males fertilize the eggs as they are deposited by females 
over logs and vegetation. The eggs are released in strings that are cemented 
together in zigzag gelatinous bands that are several inches wide and up 



272 Inland Fishes of Massachusetts 




Yellow Perch: open circles indicate known localities 
where species was not found during our post- 1969 
surveys; not all solid circles were resurveyed. 



to 7 feet long. Females may contain 2,000 to & /: £ 

157,000 eggs. There is no parental care, and egg mass 
mortality can be high as a result of predation, wind-wave action, or low- 
water conditions. Young hatch in two to three weeks, depending upon the 
water temperature. Growth is variable, but a length of 8 inches TL can be 
expected in three years. If overpopulation occurs, stunted adults seldom 
exceed 6 inches TL. Yellow Perch are diurnal carnivores, feeding on small 
aquatic insects, crustaceans, and small fishes. They are commonly found 
in schools and are active year-round. 



i.y 



distribution and abundance . Yellow Perch are distributed state- 
wide in Massachusetts, where it is a common warmwater species. 

notes . Color variants of Yellow Perch, including some with an overall blue 
wash, have been found in Massachusetts and elsewhere. Since Yellow Perch 
are active year-round, including the winter, they are popularly pursued by 
ice -anglers. 

references. Ney 1978 (review); Tsai and Gibson 1971 (fecundity); Scott 
1955, Muncy 1962, Thorpe 1977, Helfman 1979 (behavior); Jobes 1952, 
Mansueti 1964b, Brazo et al. 1975, Ney and Smith 1975 (growth); Tharratt 
1959, Siefert 1972, Keast 1977a (food); Hart 1933 (blue color); Burdick et al. 
1957 (oxygen requirements); Clady 1976 (effects of wind and temperature). 



Family and Species Accounts 273 



Walleye Introduced 

Stizostedion vitreum (Mitchill 1818) 




identification. Walleye have two well-separated dorsal fins, two poorly 
developed anal spines, two large canine teeth, and 12 to 13 anal rays. Wall- 
eye are olive-brown to gray-yellow dorsally, shading to yellow- silvery on 
the sides, and creamy- white ventrally. A dark blotch is present at the base 
of the pectoral fin and on the posterior membrane of the first dorsal fin. 
The edge of the anal fin and the tip of the lower caudal lobe are white. 

selected counts. D XIII-XIV+I,19-21; A II, 12-14; Scales 12/83-104/15; 
GR 10-13. 

size. Walleye are the largest members of the perch family. Typical adults 
range from 13 to 22 inches and 1 to 5 pounds. The Massachusetts state 
record, from Quabbin Reservoir in 1975, weighed 11 pounds. 

natural history. Walleye are a coolwater species, inhabiting both 
lakes and larger rivers. They are nocturnally active, feeding along gravel 
bars in rivers or along rocky shoals in lakes. Walleye generally prefer firm 
bottom substrates with gravel or bedrock. Similar to Yellow Perch, Walleye 
are active year-round. Walleye spawn in the spring following ice-out, when 
water temperatures approach 40°F. Males are nonterritorial, nests are not 
built, and parental care is not provided. Spawning occurs at night when one 
or two females and up to six males will move into the shallows and broad- 
cast eggs and sperm over the substrate. Females carry between 23,000 and 
615,000 eggs. Eggs deposited in crevices are protected from predation and 
hatch in 12 to 25 days, depending on water temperature. Within two weeks 
after hatching, young Walleye disperse into the shallows of open water. 
Growth is rapid, and juveniles attain a length of 3.5 to 12 inches TL by the 



274 Inland Fishes of Massachusetts 







o V w / 



Walleye: open circles indicate historic records where 
the species may no longer occur. 




/-■• 



end of their first summer in southern parts of their 
range. Male Walleye in New York mature at two or / ^ ^ 

three years of age (12 to 13 inches), while females mature 
at ages four or five (about 17 inches). The diet of Walleye shifts rapidly as 
they grow. Juveniles feed on planktonic crustaceans, gradually switching 
to aquatic insects and then to small fishes after they reach 3 inches or so. 
Adult Walleye are voracious predators that feed predominantly on other 
fishes. They can become highly cannibalistic in the absence of readily avail- 
able small fish prey. 



... ,/'■■ 



distribution and abundance . Walleye are native to a large area of 
central North America. They were first introduced into the Connecticut 
River in the early 1900s from Lake Champlain stock. From 1953 to 1960, 
stocks from western Lake Erie were introduced into several Massachusetts 
waterbodies, including Quabbin Reservoir, Lake Chauncy in Westborough, 
and Assawompsett Pond in Lakeville. In the 1980s, experimental stocking of 
a Lake Oneida strain in the Taunton River Drainage was carried out. Today, 
Walleye are found in the northern portion of the Connecticut River as well 
as in the Assawompsett Pond system of the Taunton (Nemasket) River 
Drainage. Remnant Walleye from earlier stocking programs may still be 
found in Quabbin Reservoir; however, their ability to reproduce may have 
been limited by this reservoir's characteristically acidic waters. 

notes. A subspecies of Walleye, the Blue-pike, S. v. glaucum, was formerly 
abundant in Lake Erie but is now thought to be extinct. Overfishing and 
gene mixing, coupled with the effects of accelerated eutrophication, are 
probable reasons for the loss of this subspecies. 



Family and Species Accounts 275 



references. Ney 1978, Colby et al. 1979 (review); Ryder and Kerr 1978 
(feeding); Robins 1970 (bibliography); Kelso and Ward 1972 (unexploited 
population); Campbell 1987 (Blue-pike status); Groen and Schroeder 1978 
(water level management); Nickum 1978 (culture); Serns 1978 (size limits); 
Carlander and Whitney 1961, Wolfert 1977 (age, growth); Forney 1974 (pre- 
dation); Raney and Lachner 1942, Eschmeyer 1950 (life history). 



276 Inland Fishes of Massachusetts 



Jack Family 



Carangidae 



The carangids, also known as jacks, pompano, or scad, are found worldwide 
in temperate to tropical marine waters. About 140 species are known but 
only 13 enter Massachusetts marine waters. Only one species, the Crevalle 
Jack, enters Massachusetts freshwaters. Jacks are important game and food 
fishes wherever they are found. 

references. Bigelow and Schroeder 1953 (local marine species); Laroche 
et al. 1984 (development); Smith- Vaniz 1984 (relationships). 



Crevalle Jack 

Caranx hippos (Linnaeus 1766) 



Native 




identification. Crevalle Jacks can be distinguished from all other perch- 
like Massachusetts freshwater fishes by the expanded, spinous scutes along 
the posterior straight section of the lateral line. This is the only jack found 
in local freshwater; other Caranx species, which closely resemble this spe- 
cies, are sometimes found in estuaries. Positive identification of specimens 
from estuaries may require review of the marine references at the end of 
this account. The deep-bodied juveniles are silvery with dark, vertical bands 
along the sides. Adults lack the bars, become less deep-bodied, and have a 
more rounded forehead. 

selected counts. D VII-VIII,20; A 11,17-18. 



Family and Species Accounts 277 




Crevalle Jack. 



size. Adult Crevalle Jacks grow to just over 

3 feet TL. Only small juveniles, from 3 to 5 inches 

TL, are found in Massachusetts brackish or freshwaters 



[ p\ 



K 






. ...vl3».^--.l 



J? 



natural history. Crevalle Jacks are a marine species, but, in some 
parts of their range, juveniles enter brackish to freshwater. In Massachu- 
setts, these young fishes are most commonly found between mid-June and 
mid-October. Sometimes, large numbers of juveniles enter rivers; a mass 
mortality was noted in the Slocums River, Dartmouth, in 1969. The salinity 
at the site of the die-off was almost fresh (0.5-1.4 parts per thousand); how- 
ever, these southern fishes probably died after being exposed to cold spring 
water from a nearby brook. In June 1980, we collected a juvenile (60 mm 
SL) at the head of the Westport River, which is seven miles from the river's 
mouth. When collected, this fish was healthy and had eaten a 24 mm SL 
Mummichog just before it was caught. Adult Crevalle Jacks are common 
open-water predators. 

distribution. Crevalle Jacks are found from Nova Scotia to Uruguay but 
are most common south of Cape Cod. In Massachusetts, juveniles have 
been found in the Buzzards Bay and Taunton drainages and might be ex- 
pected in the Cape Cod, Narragansett Bay, and Islands drainages. 

references. Berry and Smith -Vaniz 1977 (identification, distribution); 
Hoff 1971 (mortality, Slocums River). 



278 Inland Fishes of Massachusetts 



American Sole Family 

Achiridae 



All of the flatfishes are linked together by the asymmetrical placement of 
the eyes and their compressed bodies. In addition, flatfishes are born with 
normal eyes; that is, with an eye on each side of the head. However, during 
development and metamorphosis into the juvenile form, one eye migrates 
over the head and toward the other so that both eyes end up on one side of 
the body. The eyes almost always move to one side or the other, giving the 
flatfish either a right-eyed or left-eyed appearance that is usually character- 
istic of each family. The soles are right-eyed flounders that lack pectoral fins 
and have the blind side pelvic fin united to the anal fin. The Achiridae, con- 
sisting of about 30 species, are found only in the New World. Many of these, 
including the one Massachusetts species, enter freshwater. The true soles 
should not be confused with other flatfishes that are often marketed as 
"sole" in the United States. In New England, these latter fishes are members 
of the Pleuronectidae, of which the Winter Flounder, Pseudopleuronectes 
americanus, is marketed as "lemon sole" and the Witch Flounder, Glypto- 
cephalus cynoglossus, is sold as "gray sole." 

references. Bigelow and Schroeder 1953 (local marine species); Hensley 
and Ahlstrom 1984, Cooper and Chapleau 1998 (relationships). 



Family and Species Accounts 279 



Hogchoker Native 

Trinectes maculatus (Bloch and Schneider 1801) plate 55 




identification. Hogchokers have both eyes on the right side, lack pec- 
toral fins, and have oval bodies. Their dorsal fin reaches the tip of the snout, 
and they have a curved mouth surrounded by many fleshy tabs, or papillae. 
The eyed side is mottled brown-olive, often with 10 or so vertical lines on 
the body. The blind side is either creamy white or spotted with medium- 
sized dark dots. Several marine flatfishes may invade the upper estuary but 
should not be confused with this species because the Hogchoker is the only 
local, right- eyed flatfish that lacks pectoral fins. 

selected counts. D 50-56; A 36-42; Scales 66-75. 

size . Hogchokers are small flatfishes that seldom grow more than 6 inches 
TL. The young-of-the-year, most commonly found in freshwater, range up 
to one inch TL (25 mm SL). 

natural history. Hogchokers spend their lives on the bottom of es- 
tuaries and coastal streams. Their body shape and color provide efficient 
camouflage, and small specimens are often overlooked, even in a net, be- 
cause they look like small leaves. Hogchokers spawn from May to October 
with peak spawning occurring at maximum day length and maximum water 
temperature. The small eggs (about 1 mm) are spawned in the lower estuary 
in areas of high salinity. A large, 6-inch female may contain 50,000 eggs. 



280 Inland Fishes of Massachusetts 




Hogchoker: open circles indicate historic records 



where the species no longer occurs. 



••' 




Eggs hatch quickly, in as little as 26 to 34 hours; 
and the tiny larvae, no more than 0.8 inches TL, r i^ 

begin to move inshore toward brackish and freshwater. 
It is thought that Hogchokers use the tidal wedge for upstream transport. 
The young- of- the -year spend their first fall and winter in brackish to fresh- 
waters, where they find both protection from predators and abundant food. 
During their second summer, they move downstream but not as far as the 
spawning areas. Hogchokers mature during their third summer and live up 
to six years. Each season, adults move inshore during the winter and return 
to the spawning area in the spring. Hogchokers feed on worms, crustaceans, 
and other benthic invertebrates. 



4?\ 

-'-6;:;:... 



distribution and abundance. Massachusetts is the northernmost 
part of the range of the Hogchoker. In Massachusetts, they are most com- 
mon in the southeast, where young have been recorded from brackish and 
freshwaters of almost all the south-flowing drainages. This species is rare 
north of Cape Cod; however, L. Agassiz procured a number from the mouth 
of the Charles River in 1874. Bigelow and Schroeder (1953) noted that they 
had not seen a Hogchoker record from north of the Cape since Agassiz's re- 
port. In October 1980, we collected a juvenile in the freshwaters of Rock 
Creek, Orleans, a tributary to Cape Cod Bay; there are also recent published 
records of Hogchokers from marine waters near the Pilgrim Power Plant, 
Plymouth. 

notes. This fish's strange name originates from colonial times. Appar- 
ently, when hogs fed on discarded fishes, they had difficulty swallowing this 
fish because of its hard, rough scales. 



Family and Species Accounts 281 



references. Bigelow and Schroeder 1953 (origin of name); Dovelet al. 
1969 (life history); Koski 1978 (life history); Lawton et al. 1984 (Plymouth 
records); Martin and Drewry 1978 (review and development). 



282 Inland Fishes of Massachusetts 



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in fresh water. /. Fish Bio. 37(Supp. A) : 79-85. 

Wilson, M.V.H., and P. Veilleux. 1982. Comparative osteology and relationships of the Um- 
bridae {Pisces: Salmoniformes) . Zool. J. Linn. Soc. 76:321-352. 

Winfield, I.J., and J.S. Nelson, eds. 1991. Cyprinid fishes: systematics, biology and exploita- 
tion. Chapman & Hall (New York). 667pp. 

Winsor, M.P. 1991. Reading the shape of nature: comparative zoology at theAgassiz Mu- 
seum. Univ. Chicago Press (Chicago). 324pp. 

Wolfert, D.R. 1977. Age and growth of the walleye in Lake Erie, 1963-1968. Trans. Amer. 
Fish. Soc. 106:569-577. 

Wootton, R.J. 1976. The biology of the sticklebacks. Academic Press (London). 387pp. 

Wootton, R.J. 1984. A functional biology of sticklebacks. Univ. Cal. Press (Berkley). 265pp. 

Wootton, R.J. 1990. Ecology of teleost fishes. Fish and Fisheries Ser. No. 1. Chapman & Hall 
(London). 404pp. 

Woronecki, D.E. 1966. Some aspects of the life history of the northern rock bass, Amblo- 
plites rupestris rupestris (Rafinesque), in Quabbin Reservoir, Massachusetts. M.S. 
thesis, Univ. Mass. (Amherst). 45pp. 

Wright, D.I., and W.J. O'Brien. 1984. The development and field testing of a tactical model 
of the plantivorous feeding of white crappie {Pomoxis annularis). Ecol. Mono. 54: 
65-98. 

Wydoski, R.S., and E.L. Cooper. 1966. Maturation and fecundity of brook trout from infer- 
tile streams./. Fish. Res. Bd. Can. 23:623-649. 

Zaitzevsky, C. 1982. Frederick Law Olmsted and the Boston park system. Belknap Press. 
(Cambridge, Mass.). 262pp. 



314 Inland Fishes of Massachusetts 



Appendix 1. Indexed References 



The following references will facilitate gathering information that is beyond the 
scope of this book. For an authoritative but nontechnical overview of the world of 
fishes, we recommend Paxton and Eschmeyer (1998). The following references are 
arranged by topic. 

General Ichthyology. Bone et al. 1995, Lagler et al. 1962, Moyle and Cech 1996, Helf- 
man et al. 1997 (texts); Cailliet et al. 1986 (lab and field manual); Eschmeyer 1990, 
1998 (catalog fish genera and species); Lauder and Liem 1983, Stiassny et al. 1996 
(evolution and relationships); Marshall 1971 (life history); Moser et al. 1984 (devel- 
opment and relationships); Nelson 1994 (family review); Peden 1976 (collecting and 
preserving fishes). 

North America. Courtenay and Stauffer 1984, Fuller et al. 1999 (introduced fishes); 
Deacon et al. 1979, Ono et al. 1983, Williams et al. 1989, Williams and Miller 1990 
(rare and endangered fishes); Eschmeyer and Herald, 1983 (Pacific marine); Caven- 
der 1986 (fossils); Lee et al. 1980, Page and Burr 1991 (freshwater); Robins and Ray 
1986 (Atlantic marine); Robins et al. 1991a, 1991b (catalogs and names); Scott and 
Crossman 1973 (Canada, freshwater); Hocutt and Wiley 1986 (zoogeography); May- 
den 1992 (systematics, historical ecology); and Fishes of the Western North Atlantic 
(1948-1990+), an ongoing series now comprising nine parts and over 5,000 pages 
published as Memoirs of the Sears Foundation, Yale University, New Haven. 

Northeastern North America. Bigelowand Schroeder 1953 (Gulf of Maine); Etnier 
and Starnes 1993 (Tennessee); Everhart 1966 (Maine); Hubbs and Lagler 1964 (Great 
Lakes region); Jenkins and Burkhead 1993 (Virginia); Kendall 1908 (New England, 
annotated list); Murdy et al. 1997 (Chesapeake Bay); Scott and Scott 1988 (Canadian 
marine); Smith 1985 (New York); MacMartin 1962 (Vermont), Menhinick 1991 (North 
Carolina); Bailey 1938, Scarola 1973 (New Hampshire); Scott and Crossman 1973 
(Canada); Tracy 1906 (Rhode Island); Whitworth et al. 1968, Whitworth 1996 (Con- 
necticut); Whittier et al. 1997 (minnow), 1999, 2000, 2001 (lake fishes); Halliwell et al. 
1999, 2001 (lake fishes); Whittier 1999 (ecological classification); Daniels 1996 (fish 
scales). 

Massachusetts. Andrews 1973 (Nantucket, marine); Bigelowand Schroeder 1953, 
Clayton et al. 1978; Collette and Hartel 1988; Lawton et al. 1984 (marine); Elliott 



Appendix 1 315 



and Kushlan 1980 (ichthyoplankton, Massachusetts Bay); Halliwell 1989 (stream 
classification and habitat analysis); Hartel 1992, Cardoza et al. 1993 (introduced spe- 
cies); Hoff and Ibara 1977 (Slocums River); Kendall 1911 (Martha's Vineyard, estuar- 
ine and marine); McCabe 1942, 1943 (western Massachusetts); Storer 1839, 1867; 
and Massachusetts Wildlife, a quarterly publication of the Massachusetts Division of 
Fisheries and Wildlife, Westborough, Mass. 

Aquatic Plants and Invertebrates. Hellquist and Crow 1980, 1981, 1982, 1984; Crow 
and Hellquist 1981,1982, 1983 (aquatic plants); Pennak 1989, Smith 1991, 2001 (in- 
vertebrates); Merritt and Cummins 1996 (aquatic insects). 

Fish Larvae. Moser et al. 1984 (worldwide); Elliott and Kushlan 1980, Fahay 1983; 
Scherer 1984 (marine, North Atlantic); Lippson and Moran 1974 (lower Potomac 
River); Auer 1982 (Great Lakes). 

Ecology. Haines 1982 (acid rain); Hynes 1970 (running water); Hynes 1974, Mason 
1991 (polluted water); Matthews and Heins 1987 (streams); Reid and Wood 1976, 
Wootton 1990 (general). 

Fisheries and Fish Management. Carlander 1969, 1977, Lagler 1956, Kendall 1978, 
Schreck and Moyle 1990, Murphy and Willis 1996. 

Professional Journals. The majority of the journals that are referenced in this book 
are technical and can be found only in academic libraries. A few of the more impor- 
tant are: Copeia, the quarterly journal of The American Society of Ichthyologists and 
Herpetologists; Transactions of the American Fisheries Society, The North American 
Journal of Fish Management, and The Progressive Fish-Culturist, all published by the 
American Fisheries Society; Environmental Biology of Fishes, Kluwer Academic Pub- 
lishers Group, the Netherlands; and Canadian Journal of Fisheries and Aquatic Sci- 
ences, National Research Council, Canada. 



316 Inland Fishes of Massachusetts 



Appendix 2. Distribution Table 
of Massachusetts Inland Fishes 



Appendix 2 317 



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320 Inland Fishes of Massachusetts 



Appendix 3. Glossary 



Acid precipitation Rain or 
snow with a characteristically 
lowpH (acidic). 

Adipose fin, dorsal A small 
fleshy fin along the dorsal 
midline before the caudal fin. 

Aestivation A form of in- 
activity similar in some ways 
to hibernation. 

Ammocoete A larval 
lamprey. 

Amphipods Small shrimp - 
like crustaceans. 

Anadromous Fishes that live 
in marine waters and migrate 
into freshwater to spawn; see 
catadromous. 

Anal fin The ventral median 
fin, located behind the anus. 

Anoxic Lacking oxygen. 

Anus The vent or terminal 
opening of the intestine. 

Asymmetrical Structures 
shaped differently on left or 
right planes of body (not 
symmetrical). 

Axillary projection (or scale) 

An elongate scale at the base 
of the pelvic fin. 

Barbel Fleshy projection 
containing sensory organs; 
sometimes long and pointed 
(as in catfish) and located on 
the snout, jaws, chin, or lips. 



Benthic Associated with the 
the substrate or bottom. 

Buccal cavity The mouth or 
oral cavity. 

Catadromous Fishes that 
live in freshwater and mi- 
grate to salt water to spawn; 
see anadromous. 

Caudal fin The posterior 
median fin, often called the 
tail fin. 

Caudal peduncle The part of 
the body that supports the 
tail, starting at the end of the 
anal fin. 

Cheek The side of the head 
below and slightly behind the 
eye. 

Chironomids Nonbiting 
midges (flies, Chironomidae). 

Cladocerans A group of 
small, flea-like crustaceans 
(zooplankton) . 

Cloacal appendage Tissue 
adjacent to the cloaca, en- 
larged, often pointed. 

Compressed Laterally com- 
pressed, flattened side to side, 
as in the body of many fishes. 

Copepods A group of small 
shrimplike crustaceans 
(zooplankton). 

Crepuscular Most active at 
dawn or dusk. 



Crustaceans A large group 
of invertebrates, including 
crabs, shrimp, crayfish, and 
copepods. 

Ctenoid scale A scale with 
spines on posterior edge. 

Cycloid scale A scale with a 
smooth edge. 

Deep-bodied A body rela- 
tively deep in relation to 
length. 

Depauperate Refers to a 
fauna comprised of few 
species. 

Diadromous Fishes that mi- 
grate between saltwater and 
freshwater (anadromous and 
catadromous). 

Diatoms A group of pelagic 
microscopic algae. 

Dorsal fin The dorsal me- 
dian fin that may be divided 
into two parts and supported 
by soft rays or a combination 
of spiny and soft rays. 

Endemic Native and re- 
stricted to an area. 

Exotic Not native, intro- 
duced to an area by humans. 

Extinction The complete 
and permanent loss of a 
taxon from its total range. 

Extirpated Refers to a taxon 
(species, subspecies, etc.) 



Appendix 3 321 



lost from parts of its total 
range. 

Falcate Usually referring to 
fins that are notably concave 
or sickle-shaped on the trail- 
ing edge. 

Fauna Refers to animals that 
occur in a given region. 

Fecundity The reproductive 
potential of a female based 
on the number of eggs pro- 
duced annually. 

Filamentous Pertaining to 
long, threadlike structures. 

Fin rays Structures that sup- 
port the fin; always paired, 
usually segmented and 
branched but may be spine- 
like in some fishes. 

Fin spines Bony structures 
that support fins; usually 
pointed and stiff, never seg- 
mented or branched. 

Frenum A band of tissue 
linking the upper lip to the 
snout. 

Gas bladder See swim 
bladder. 

Gill filaments The posterior 
projections of the gill arch 
used in respiration. 

Gill rakers The anterior pro- 
jection of the gill arch (see 
Figure 8); often used to filter 
or process food. 

Gonads The reproductive 
organs: ovaries or testes. 

Gular plate A bony element 
on the ventral part of the 
head as found in Bowfin. 

Heterocercal A caudal or tail 
fin in which the vertebrae ex- 
tend into and support the 
upper tail lobe. 



Homocercal Asymmetrical 
tail in which the vertebrae 
are not turned into the upper 
lobe. 

Interorbit (interorbital) The 

area between the eyes, mea- 
sured betwen the bony rims 
of the eyes. 

Introduced (alien, exotic, 
or non-native) Transported 
by humans into an area in 
which it did not historically 
occur. 

Larva The early life stage of 
a fish, usually with a different 
body shape and behavior. 

Lateral line Pores along the 
body where neuro-receptors 
penetrate scales, bony ele- 
ments, or skin. 

Leptocephalus The larval 
stage of eel-like fishes. 

Maxilla The posterior up- 
per jaw bone in fishes (see 
Figure 8). 

Midges See Chironomids. 

Milt Sperm and associated 
fluid, often cloudy when re- 
leased into water. 

Nape The part of the back 
from the posterior end of the 
head to the origin of the dor- 
sal fin. 

Nasal Pertaining to the nose. 

Native Species known to oc- 
cur naturally in an area, not 
introduced by humans. 

Nocturnal Pertaining to ac- 
tivity at night. 

Opercle Part of a major set 
of bones on the side of the 
head that protects the gills, 
sometimes called a gill cover 
or operculum (see Figure 8). 



Oral disc The mouth struc- 
ture of a lamprey. Larval lam- 
prey have a fleshy oral hood 
covering the area. 

Oral jaws The front or typi- 
cal jaws in contrast to the in- 
ternal or pharyngeal jaws. 
Consists of the premaxilla, 
maxilla, and dentary. 

Origin Used in reference to 
structures; for example, the 
beginning or anterior-most 
starting point of a fin. 

Papilla Small fleshy dermal 
structures that are often 
knoblike. 

Parr Juvenile salmon or 
trout, often with dark vertical 
bars on body (parr marks). 

Pectoral fins Paired fins as- 
sociated with pectoral girdle 
or "shoulder bones," can be 
low or high on body. 

Pelvic axillary process A 

modified scale located in the 
axil of the pelvic fin. 

Pelvic fins A set of paired 
fins (also see pectoral fin) 
that are highly variable in 
shape and position but usu- 
ally found ventrally, posterior 
to pectoral fins. 

Peritoneum The lining of 
the abdominal cavity. 

Pharyngeal Pertaining to the 
throat in the area of the bran- 
chial basket or gill arches. 

Pleat Structure of soft tissue 
usually found in rows, ridges, 
or grooves as on the lips of 
suckers. 

Predorsal The area anterior 
to the dorsal fin origin. 

Premaxilla The anterior 
bones of the upper oral jaw 
(see Figure 8). 



322 Inland Fishes of Massachusetts 



Preoperculum Part of a ma- 
jor set of bones on the side of 
the head that protects the 
gills, sometimes called gill 
cover. 

Primary Refers to the 
groups of fishes that live only 
in freshwater. 

Procurrent rays Small rays 
anterior to median fins, most 
evident before the ventral 
and dorsal edge of the cau- 
dal fin. 

Protrusiblejaw Jaw in 

which the structure of bones 
and muscles allow the jaw to 
protrude from the snout. 

Pyloric caeca Out-pocket- 
ings along the digestive tract. 

Redd A nest created in 
gravel substrate by salmon 
and trout. 

Refugium An area where a 
species may survive adverse 
conditions. 

Ripe Gravid or containing 
developed eggs or sperm; 
ready to spawn. 



Salinity The amount or con- 
centration of salt in water, 
usually expressed in parts 
per thousand. 

Secondary Refers to a group 
of fishes that regularly move 
between freshwater and salt- 
water. 

Serrated A structure with a 
sawlike edge; usually a fin 
spine. 

SL Standard Length, a 
method of measuring fishes 
(see Figure 8). 

Slab-sided Describes a fish 
with a flat-sided body, which 
is compressed laterally. 

Snout The area of the head 
anterior to the eyes. 

Soft rays Supporting ele- 
ments in all fins, usually 
branched and always seg- 
mented unlike fin spines. 
Also see fin rays. 

Spiral valve Specialized 
internal structures in the 
intestine. 



Swim bladder A bladderlike 
structure found dorsally in 
the abdominal cavity; often 
filled with gas (sometimes 
called gas bladder) and used 
for buoyancy control, respi- 
ration, and gas exchange. 

TL Total Length, a method 
of measuring fishes (see Fig- 
ure 8). 

Vascularized Tissue con- 
taining many small blood 
vessels. 

Viscera The internal organs, 
such as heart, stomach, liver, 
and kidneys. 

Vomer Bone found along 
midline of the roof of the 
mouth; often with teeth. 

Zoogeography The study of 
the geographical distribution 
of animals. 

Zooplankton The portion of 
the plankton composed of 
small animals rather than 
plants. 



Appendix 3 323 



Taxonomic Index 



Ablennes, 195 
Acanthopterygii, 4, 6 
Achiridae, 5, 51, 279 
Acipenser, 67-70 
Acipenseridae, 4, 66 

key to, 66 
Actinopterygii, 4, 6 
aculeatus, Gasterosteus, 223 
acus, Tylosurus, 196 
aenaeus, Myoxocephalus, 

232 
aestivalis, Alosa, 80 
affinis, Gambusia, 34, 198- 

199 
Agnatha, 4 
Agujon, 196 
Alewife, 10, 17, 24, 82 
Alewife Floater, 86 
Alosa, 80-86 
Ambloplites, 246 
Ameiurus, 146-151 
americana, Morone, 236 
americanus, Esox, 159 
Amia, 4, 71 
Amiidae, 4, 71 
Ammocoetes, 60 
Anchoa, 89 
Anchovy 

Bay, 38, 89 
Anguilla, 4, 75 
Anguillidae, 4, 74 
Anguilliformes, 4, 74 
annularis, Pomoxis, 262 
Anodonta, 86 
Apeltes, 221 
Aphredoderidae, 186 
appendix, Lampetra, 62 
Ariidae, 143 
Astronotus, 34 
Atherinidae, 211 



Atheriniformes, 4 
atherinoides, Notropis, 34, 

92 
Atherinomorpha, 4, 6 
Atherinopsidae, 4, 211 

key to, 211 
atratulus, Rhinichthys, 124 
atromaculatus, Semotilus, 

130 
auratus, Carassius, 100 
auritus, Lepomis, 250 

B 

bairdi, Cottus, 233 
Bass 

key to, 235, 242 

Largemouth, 23, 25, 260 

Rock, 246 

Smallmouth, 258 

Striped, 9-10, 18-19, 238 
batrachus, Clarias, 34, 143 
Belonidae, 4, 195 
beryllina, Menidia, 212 
bifrenatus, Notropis, 112 
Bluegill, 23, 256 
Bowfin, 4, 71 

brachypomus, Piractus, 34 
brevirostrum, Acipenser, 67 
Brevoortia, 78 
Bullhead 

Black, 145, 149 

Brown, 22, 23, 149 

Yellow, 148 
Bullhead Catfishes, 4, 143 

key to, 144 
Burbot, 22, 37, 190 



calva, Amia, 71 
Capelin, 168 
Carangidae, 5, 277 



Caranx, 277 
Carassius, 100 
Carp 

Common, 104 

Grass, 34, 92, 95 

key to, 93 

Leather, 104 

Mirror, 104 
carpio, Cyprinus, 104 
cataractae, Rhinichthys, 

126 
Catfish 

Channel, 151 

Upside-down, 34 

Walking, 34, 143 

White, 146 
Catfishes, 4, 143 

key to, 144 
Catostomidae, 4, 135 

key to, 136 
Catostomus, 137-140 
catostomus, Catostomus, 

137 
catus, Ameiurus, 146 
Centrarchidae, 5, 241 

key to, 242 
cepedianum, Dorosoma, 86 
Cephalaspidomorphi, 4 
cephalus, Mugil, 217 
Channa, 34 
Char(s) 

Brook, see trout, brook 

Lake, see trout, lake 
Chrosomus, 119 
Chub 

Creek, 37, 130 

key to, 93 

Lake, 22, 37, 102 
Chubsucker 

Creek, 141 

Lake, 142 



325 



Cichlasoma, 34 
Cichlid, Midas, 34 
citrinellum, Cichlasoma, 34 
Clarias, 143 
Clupea, 78 
Clupeidae, 4, 78 

key to, 79 
Clupeiformes, 4, 89 
Clupeomorpha, 4, 6 
Coelocanth, 31 
Codfishes, 189 
cognatus, Cottus, 232 
Coho, 171 
Colossoma, 34 
commersoni, Catostomus, 

139 
Coregonidae, 171 
Coregoninae, 171 
cornutus, Luxilus, 108 
corporalis, Semotilus, 132 
Cottidae, 5, 231 
Cottus, 232 
Couesius, 102 
Crappie 

Black, 264 

key to, 243 

White, 262 
crysoleucas, Notemigonus, 

110 
Ctenopharyngodon, 92, 95 
curema, Mugil, 216 
Cusk, 189 

cyanellus, Lepomis, 252 
Cyprinella, 34 
Cyprinidae, 4, 92 

key to, 93 
Cyprinodon, 201 
Cyprinodontidae, 5, 198 

key to, 199 
Cyprinus, 104 

D 

Dace 

Blacknose, 15, 22, 124 

Finescale, 118 

key to, 93 

Longnose, 12, 15, 22, 38, 
126 

Northern Redbelly, 22, 
37, 118 
Darter 

Johnny, 271 



key to, 266 

Swamp, 12, 14, 23, 25, 37, 
268 

Tessellated, 12, 15, 23, 
270 
diaphanus, Fundulus, 203 
Diplurus, 31-32 
Dipnoi, 6 

dolomieu, Micropterus, 258 
Dorosoma, 86 

E 

Eel 

American, 9, 23, 33, 75 

European, 74 
Eels, Freshwater, 4, 74 
Elopomorpha, 4, 6 
Engraulidae, 4, 89 
Engraulis, 89 
Enneacanthus, 248 
eos, Phoxinus, 118 
Erimyzon, 141 
erythrophthalmus, Scardi- 

nius, 128 
Esocidae, 4, 157 

key to, 158 
Esociformes, 4 
Esox, 159-164 
Etheostoma, 268-272 
Etheostomatini, 266 
eurystole, Engraulis, 89 
Exoglossum, 34, 93-94 



Fallfish, 8, 15, 23, 38, 132 
flauescens, Perca, 272 
fontinalis, Salvelinus, 181 
Fundulidae, 5, 198 

key to, 199 
Fundulus, 198, 203-208 
fuscus, Syngnathus, 59, 229 
fusiforme, Etheostoma, 268 



Gadidae, 4, 189 

key to, 189 
gairdneri, Salmo, 175 
Gambusia, 34, 198 
Gar, 34 
Gasterosteidae, 5, 219 

key to, 219 
Gasterosteiformes, 5 



Gasterosteus, 223-226 
gibbosus, Lepomis, 254 
glaucum, Stizostedion v., 

275 
Gnathostomata, 4, 6 
Goldfish, 100 
Grubby, 232 
gyrinus, Noturus, 153 

H 

Haddock, 189 
Hake, 189 

harengus, Clupea, 78 
hepsetus, Anchoa, 89 
Herring, 4, 78 

Atlantic, 78 

Blueback, 17-19, 80 

key to, 79 
heteroclitus, Fundulus, 

205 
hians, Ablennes, 195 
hippos, Caranx, 277 
Hogchoker, 280 
hudsonius, Notropis, 114 
Hybognathus, 106 
Hypentelium, 34, 135 

I 

Ictaluridae, 4, 143 

key to, 144 
Ictalurus, 151 
idella, Ctenopharyngodon, 

34, 92, 95 
implicata, Anodonta, 86 
insignis, Noturus, 155 
insulae, Etheostoma f., 269 

J 

Jack, Crevalle, 277 

K 

Killifish 

Banded, 203 

Rainwater, 203, 36, 209 

Spotfin, 207 

Striped, 198, 200 

Waccamaw, 204 
Killifishes, 5, 198 

key to, 199 
kisutch, Oncorhynchus, 171 
kokanee, 171 
krameri, Umbra, 165 



326 Inland Fishes of Massachusetts 



lacerum, Moxostoma, 135 
Lampetra, 62 
Lamprey, 4, 60 

American Brook, 22, 37, 
62 

key to, 61 

Sea, 10, 64 
Latimeria, 31 
Lepisosteus, 34 
Lepomis, 250-258 
limi, Umbra, 165 
Lota, 190 
/ota, Lota, 190 
Lucania, 209 
luciae, Fundulus, 207 
Luciopercini, 266 
lucius, Esox, 161 
lutrensis, Cyprinella, 34 
Luxilus, 108 

M 

macrochirus, Lepomis, 256 
macrolepidotus, Fundulus 

h., 206 
macropomum, Colossoma, 

34 
maculatus, Trinectes, 280 
Madtom 

key to, 145 

Margined, 155 

Tadpole, 153 
majalis, Fundulus, 198, 200 
marina, Strongylura, 195 
marinus, Petromyzon, 64 
masquinongy, Esox, 162 

hybrid, 158, 162 
maxillingua, Exoglossum, 

34,92-94 
mediocris, Alosa, 78-79 
melas, Ameiurus, 145, 149 
Menhaden, Atlantic, 78 
Menidia, 212-215 
menidia, Menidia, 214 
metae-gadi, Etheostomaf., 

269 
Microgadus, 192 
micropeltes, Channa, 34 
Micropterus, 258-261 
Minnow 

Bluntnose, 14, 33, 120 

Cutlips, 34, 92-94 



Eastern Silvery, 22, 37, 
106 

Fathead, 122 

key to, 93 

Sheepshead, 201 
mitchilli, Anchoa, 89 
mordax, Osmerus, 168 
Morone, 235-240 
Moronidae, 5, 235 

key to, 235 
Mosquitofish, 34, 198-199 
Moxostoma, 135 
Mudminnow, Central, 165 
Mugil, 216-218 
Mugilidae, 4, 216 

key to, 216 
Mugilomorpha, 4, 6 
Mullet 

key to, 216 

Striped, 216-217 

White, 217 
Mummichog, 36, 205 
Muskellunge, Tiger, 158, 

161-162 
my kiss, Oncorhynchus, 

174 
Myoxocephalus, 231-232 

N 

namaycush, Salvelinus, 183 

nannomyzon, C. catosto- 

mus, 138 
natalis, Ameiurus, 148 
nattered, Pygocentrys, 34 
nebulosus, Ameiurus, 149 
Needlefish, 4, 195 
Atlantic, 195 
Flat, 195 
neogaeus, Phoxinus, 118 
Neognathi, 4, 6 
nerka, Oncorhynchus, 171 
niger, Esox, 163 
nigricans, Hypentelium, 34, 

135 
nigromaculatus, Pomoxis, 

264 
nigrum, Etheostoma, 271 
notatus, Pimephales, 120 
Notemigonus, 110 
Notropis, 112-117 
Noturus, 153-156 
nuchalis, Hybognathus, 108 



O 

obesus, Enneacanthus, 248 
oblongus, Erimyzon, 141 
occidentalis, Pungitius, 

228 
ocellatus, Astronotus, 34 
oculatus, Lepisosteus, 34 
olmstedi, Etheostoma, 270 
omiscomaycus, Percopsis, 

186 
Oncorhynchus, 171 
Oscar, 34 
Osmeridae, 4, 168 
Osmerus, 168 
Ostariophysi, 4, 6 
oxyrinchus, Acipenser, 69 

P 

Pacu, 34 

Paracanthopterygii, 4, 6 
parva, Lucania, 209 
P<?rc<2, 272 
Perch 

key to, 235, 266 

White, 10, 236 

Yellow, 10, 23, 25, 272 
Percichthyidae, see 

Moronidae 
Percidae, 5, 266 

key to, 266 
Perciformes, 5 
Percini, 266 
Percomorpha, 5, 6 
Percopsidae, 4, 186 
Percopsis, 186 
Petromyzon, 64 
Petromyzontidae, 4, 60 

key to, 60 
Phoxinus, 118 
Pickerel 

Bulldog, 160 

Chain, 8, 23, 163 

key to, 158 

Mud, 160 

Redfin, 23, 25, 37, 159 
Pike 

key to, 158 

Northern, 161 
Pimephales, 120-124 
Pipefish, Northern, 12, 229 
Piractus, 34 
Piranah, Red, 34 



Taxonomic Index 327 



Pirapatinga, 34 
Pirate Perch, 186 
Pleuronectiformes, 5, 7 
plumbeus, Couesius, 102 
Poeciliidae, 198 
Pomoxis, 262-265 
promelas, Pimephales, 122 
Protocanthopterygii, 4, 6 
pseudoharengus, Alosa, 82 
Pumpkinseed, 10, 23, 25, 

254 
punctatus, Ictalurus, 151 
Pungitius, 227 
pungitius, Pungitius, 227 
Pupfish, 5, 199 

key to, 199 
Pygocentrus, 34 



quadracus, Apeltes, 221 

R 

Redfieldius, 31-32 
regius, Hybognathus, 106 
Rhinichthys, 124-128 
rostrata, Anguilla, 75 
Rudd, 128 
rupestris, Ambloplites, 246 



sfl/ar, Salmo, 9, 176 
Sfl/mo, 176-181 
salmoides, Micropterus, 260 
Salmon, 4, 171 

Atlantic, 9, 17-23, 33, 176 

Coho, 171 

Sockeye, 171 
Salmonidae, 4, 171 

key to, 172 
Salmons, Pacific, 171-172 
Salvelinus, 181-185 
sapidissima, Alosa, 84 
Sarcopterygii, 6 
saxatilis, Morone, 238 
Scardinius, 128 
Scorpaeniformes, 5 
Sculpin, 

Deepwater, 231 



Mottled, 233 

Slimy, 12, 22, 23, 37, 232 
Semionotus, 31-32 
Semotilus, 130-134 
Shad 

American, 9, 16-19, 21, 84 

Gizzard, 86 

Hickory, 78-79 

key to, 79 
Shiner 

Bridle, 14, 22, 112 

Common, 22-23, 108 

Emerald, 34, 92, 98 

Golden, 23, 110 

key to, 93 

Mimic, 116 

Red, 34 

Spottail, 114 
Silverside 

Atlantic, 214 

Brook, 211 

Inland, 36, 38, 212 
Silversides, 4, 211 

key to, 211 
Smelt, Rainbow, 10, 16, 168 
Snakehead, 34 
Sole, 279 
Stickleback 

Blackspotted, 225 

Fourspine, 221 

key to, 219 

Ninespine, 227 

Threespine, 22, 223 
Stizostedion, 274 
Strongylura, 195 
Sturgeon, 4, 66 

Atlantic, 22, 69 

key to, 66 

Shortnose, 14, 22, 67 
sucetta, Erimyzon, 142 
Sucker 

key to, 136 

Longnose, 22, 37, 137 

Northern Hog, 34, 135 

White, 22, 139 
Sunfish 

Banded, 12, 23, 25, 37, 
248 



Green, 252 

key to, 242 

Longear, 251 

Redbreast, 23, 250 
Syngnathidae, 5, 229 
Syngnathiformes, 5 
Syngnathus, 229 
Synodontis, 34 



Tambaqui, 34 
Teleostei, 4, 6 
thompsoni, Myoxo- 

cephalus, 231 
Tomcod, Atlantic, 9, 192 
tomcod, Microgadus, 192 
Trinectes, 280 
Trout 

Brook, 8, 22, 181 

Brown, 179 

key to, 172 

Lake, 23, 183 

LochLeven, 180 

Rainbow, 174 
Trout-perch, 4, 33, 37, 186 
trutta, Salmo, 179 
Tylosurus, 196 
tyrannus, Brevoortia, 78 

U 

Umbra, 4, 165 
Umbridae, 4, 165 



variegatus, Cyprinodon, 

201 
vitreum, Stizostedion, 274 
volucellus, Notropis, 116 

W 

waccamensis, Fundulus, 

204 
Walleye, 274 
wheatlandi, Gasterosteus, 

225 
Whitings, 189 



328 Inland Fishes of Massachusetts 



HUDSON 




VERMONT 




NEW HAMPS 



CON 




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Falls, 



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Petersham 




Princetc] 



Amherst 
North a mp tori^ 



hi 




Quablnn 
Reservoir 



West field 



SJJNT 

• Spring fi/eld 




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HOUS ATONIC 



IVER 




CONNECTICUT 




THAMES 



RIV 



CJ?* 



Long Island Sound 




Montauk 




Wellfleet 



NANTUCKET